Gymnospermae.—Trees or shrubs; leaves vary considerably in size and form. Flowers unisexual, except in a few cases (Gnetales) without a perianth. Monoecious or dioecious. Ovules naked, rarely without carpellary leaves, usually borne on carpophylls, which assume various forms. The single megaspore enclosed in the nucellus is filled with tissue (prothallus) before fertilization, and contains two or more archegonia, consisting usually of a large egg-cell and a small neck, rarely of an egg-cell only and no neck (GnetumandWelwitschia). Microspore spherical or oval, with or without a bladder-like extension of the exine, containing a prothallus of two or more cells, one of which produces two non-motile or motile male cells. Cotyledons two or several. Secondary xylem and phloem produced by a single cambium, or by successive cambial zones; no true vessels (except in the Gnetales) in the wood, and no companion-cells in the phloem.I.Pteridospermae(seePalaeobotany,Palaeozoic).II.Cycadophyta.A. Cycadales (recent and extinct).B. Bennettitales (see Palaeobotany:Mesozoic).III.Cordaitales(seePalaeobotany:Palaeozoic).IV.Ginkgoales(recent and extinct).V.Coniferales.A. Taxaceae.B. Pinaceae.There is no doubt that the result of recent research and of work now in progress will be to modify considerably the grouping of the conifers. The familyAraucarieae, represented byAraucariaandAgathis, should perhaps be separated as a special class and a rearrangement of other genera more in accord with a natural system of classification will soon be possible; but for the present its twofold subdivision may be retained.VI.Gnetales.A. Ephedroideae.B. Gnetoideae.C. Welwitschioideae (Tumboideae).Cycadophyta.—A.Cycadales.—Stems tuberous or columnar, not infrequently branched, rarely epiphytic (Peruvian species ofZamia); fronds pinnate, bi-pinnate in the Australian genusBowenia. Dioecious; flowers in the form of cones, except the female flowers ofCycas, which consist of a rosette of leaf-like carpels at the apex of the stem. Seeds albuminous, with one integument; the single embryo, usually bearing two partially fused cotyledons, is attached to a long tangled suspensor. Stems and roots increase in diameter by secondary thickening, the secondary wood being produced by one cambium or developed from successive cambium-rings.Fig. 1.—Stem ofCycas.F, foliage-leaf bases;S, scale-leaf bases.Fig. 2.—Cycas siamensis.The cycads constitute a homogeneous group of a few living members confined to tropical and sub-tropical regions. As a fairly typical and well-known example of the Cycadaceae, a species of the genusCycas(e.g.C. circinalis,C. revoluta, &c.) is briefly described. The stout columnar stem may reach a height of 20 metres, and a diameter of half a metre; it remains either unbranched or divides near the summit into several short and thick branches, each branch terminating in a crown of long pinnate leaves. The surface of the stem is covered with rhomboidal areas, which represent the persistent bases of foliage- and scale-leaves. In some species ofCycasthere is a well-defined alternation of transverse zones on the stem, consisting of larger areas representing foliage-leaf bases, and similar but smaller areas formed by the bases of scale-leaves (FandS, fig. 1). The scale-leaves clothing the terminal bud are linear-lanceolate in form, and of a brown or yellow colour; they are pushed aside as the stem-axis elongates and becomes shrivelled, finally falling off, leaving projecting bases which are eventually cut off at a still lower level. Similarly, the dead fronds fall off, leaving a ragged petiole, which is afterwards separated from the stem by an absciss-layer a short distance above the base. In some species ofCycasthe leaf-bases do not persist as a permanent covering to the stem, but the surfaceis covered with a wrinkled bark, as inCycas siamensis, which has a stem of unusual form (fig. 2). Small tuberous shoots, comparable on a large scale with the bulbils ofLycopodium Selago, are occasionally produced in the axils of some of the persistent leaf-bases; these are characteristic of sickly plants, and serve as a means of vegetative reproduction. In the genusCycasthe female flower is peculiar among cycads in consisting of a terminal crown of separate leaf-like carpels several inches in length; the apical portion of each carpellary leaf may be broadly triangular in form, and deeply dissected on the margins into narrow woolly appendages like rudimentary pinnae. From the lower part of a carpel are produced several laterally placed ovules, which become bright red or orange on ripening; the bright fleshy seeds, which in some species are as large as a goose’s egg, and the tawny spreading carpels produce a pleasing combination of colour in the midst of the long dark-green fronds, which curve gracefully upwards and outwards from the summit of the columnar stem. InCycasthe stem apex, after producing a cluster of carpellary leaves, continues to elongate and produces more bud-scales, which are afterwards pushed aside as a fresh crown of fronds is developed. The young leaves ofCycasconsist of a straight rachis bearing numerous linear pinnae, traversed by a single midrib; the pinnae are circinately coiled like the leaf of a fern (fig. 3). The male flower ofCycasconforms to the type of structure characteristic of the cycads, and consists of a long cone of numerous sporophylls bearing many oval pollen-sacs on their lower faces. The type described serves as a convenient representative of its class. There are eight other living genera, which may be classified as follows:—Fig. 3.—Cycas.Young Frond.Classification.—A.Cycadeae.—Characterized by (a) the alternation of scale- and foliage-leaves (fig. 1) on the branched or unbranched stem; (b) the growth of the main stem through the female flower; (c) the presence of a prominent single vein in the linear pinnae; (d) the structure of the female flower, which is peculiar in not having the form of a cone, but consists of numerous independent carpels, each of which bears two or more lateral ovules. Represented by a single genus,Cycas. (Tropical Asia, Australia, &c.).B.Zamieae.—The stem does not grow through the female flower; both male and female flowers are in the form of cones. (a)Stangerieae.—Characterized by the fern-like venation of the pinnae, which have a prominent midrib, giving off at a wide angle simple or forked and occasionally anastomosing lateral veins. A single genus,Stangeria, confined to South Africa, (b)Euzamieae.—The pinnae are traversed by several parallel veins.Bowenia, an Australian cycad, is peculiar in having bi-pinnate fronds (fig. 5). The various genera are distinguished from one another by the shape and manner of attachment of the pinnae, the form of the carpellary scales, and to some extent by anatomical characters.Encephalartos(South and Tropical Africa).—Large cones; the carpellary scales terminate in a peltate distal expansion.Macrozamia(Australia).—Similar toEncephalartosexcept in the presence of a spinous projection from the swollen distal end of the carpels.Zamia(South America, Florida, &c.).—Stem short and often divided into several columnar branches. Each carpel terminates in a peltate head.Ceratozamia(Mexico).—Similar in habit toMacrozamia, but distinguished by the presence of two horn-like spinous processes on the apex of the carpels.Microcycas(Cuba).—LikeZamia, except that the ends of the stamens are flat, while the apices of the carpels are peltate.Dioon(Mexico) (fig. 4).—Characterized by the woolly scale-leaves and carpels; the latter terminate in a thick laminar expansion of triangular form, bearing two placental cushions, on which the ovules are situated.Bowenia(Australia).—Bi-pinnate fronds; stem short and tuberous (fig. 5).From a photograph of a plant in Peradeniya Gardens, Ceylon, by Professor R. H. Yapp.Fig. 5.—Bowenia spectabilis: frond.Fig. 4.—Dioon edule.Fig. 6.—Macrozamia heteromera.A, part of frond;B, single pinna.The stems of cycads are often described as unbranched; it is true that in comparison with conifers, in which the numerous branches, springing from the main stem, give a characteristic form to the tree, the tuberous or columnar stem of the CycadaceaeStem and leaf.constitutes a striking distinguishing feature. Branching, however, occurs not infrequently: inCycasthe tall stem often produces several candelabra-like arms; inZamiathe main axis may break up near the base into several cylindrical branches; in species ofDioon(fig. 4) lateral branches are occasionally produced. The South AfricanEncephalartosfrequently produces several branches. Probably the oldest example of this genus in cultivation is in the Botanic Garden of Amsterdam, its age is considered by Professor de Vries to be about two thousand years: although an accurate determination of age is impossible, there is no doubt that many cycads grow very slowly and are remarkable for longevity. The thick armour of petiole-bases enveloping the stem is a characteristic Cycadean feature; inCycasthe alternation of scale-leaves and fronds is more clearly shown than in other cycads; inEncephalartos,Dioon, &c., the persistent scale-leaf bases are almost equal in size to those of the foliage-leaves, and there is no regular alternation of zones such as characterizes some species ofCycas. Another type of stem is illustrated byStangeriaandZamia, also by a few forms ofCycas(fig. 2), in which the fronds fall off completely, leaving a comparatively smooth stem. TheCyastype of frond, except as regards the presence of a midrib in each pinna, characterizes the cycads generally, exceptBoweniaandStangeria. In the monotypic genusBoweniathe large fronds, borne singly on the short and thick stem, are bi-pinnate (fig. 5); the segments, which are broadly ovate or rhomboidal, have several forked spreading veins, and resemble the large pinnules of some species ofAdiantum. InStangeria, also a genus represented by one species (S. paradoxaof South Africa), the long and comparatively broad pinnae, with an entire or irregularly incised margin, are very fern-like, a circumstance which led Kunze to describe the plant in 1835 as a species of the fernLomaria. In rare cases the pinnae of cycads are lobed or branched: inDioon spinulosum(Central America) the margin of the segments bears numerous spinous processes; in some species ofEncephalartos,e.g.E. horridus, the lamina is deeply lobed; and in a species of the Australian genusMacrozamia,M. heteromera, the narrow pinnae are dichotomously branched almost to the base (fig. 6), and resemble the frond of some species of the fernSchizaea, or the fossil genusBaiera(Ginkgoales). An interesting species ofCycas,C. Micholitzii, has recently been described by Sir William Thiselton-Dyer from Annam, where it was collected by one of Messrs Sanders & Son’s collectors, in which the pinnae instead of being of the usual simple type aredichotomously branched as inMacrozamia heteromera. InCeratozamiathe broad petiole-base is characterized by the presence of two lateral spinous processes, suggesting stipular appendages, comparable, on a reduced scale, with the large stipules of the Marattiaceae among Ferns. The vernation varies in different genera; inCycasthe rachis is straight and the pinnae circinately coiled (fig. 3); inEncephalartos,Dioon, &c., both rachis and segments are straight; inZamiathe rachis is bent or slightly coiled, bearing straight pinnae. The young leaves arise on the stem-apex as conical protuberances with winged borders on which the pinnae appear as rounded humps, usually in basipetal order; the scale-leaves in their young condition resemble fronds, but the lamina remains undeveloped. A feature of interest in connexion with the phylogeny of cycads is the presence of long hairs clothing the scale-leaves, and forming a cap on the summit of the stem-apex or attached to the bases of petioles; on some fossil cycadean plants these outgrowths have the form of scales, and are identical in structure with the ramenta (paleae) of the majority of ferns.The male flowers of cycads are constructed on a uniform plan, and in all cases consist of an axis bearing crowded, spirally disposed sporophylls. These are often wedge-shaped and angular; in some cases they consist of a short, thickFlower.stalk, terminating in a peltate expansion, or prolonged upwards in the form of a triangular lamina. The sporangia (pollen-sacs), which occur on the under-side of the stamens, are often arranged in more or less definite groups or sori, interspersed with hairs (paraphyses); dehiscence takes place along a line marked out by the occurrence of smaller and thinner-walled cells bounded by larger and thicker-walled elements, which form a fairly prominent cap-like “annulus” near the apex of the sporangium, not unlike the annulus characteristic of the Schizaeaceae among ferns. The sporangial wall, consisting of several layers of cells, encloses a cavity containing numerous oval spores (pollen-grains). In structure a cycadean sporangium recalls those of certain ferns (Marattiaceae, Osmundaceae and Schizaeaceae), but in the development of the spores there are certain peculiarities not met with among the Vascular Cryptogams. With the exception ofCycas, the female flowers are also in the form of cones, bearing numerous carpellary scales. InCycas revolutaandC. circinaliseach leaf-like carpel may produce several laterally attached ovules, but inC. Normanbyanathe carpel is shorter and the ovules are reduced to two; this latter type brings us nearer to the carpels ofDioon, in which the flower has the form of a cone, and the distal end of the carpels is longer and more leaf-like than in the other genera of theZamieae, which are characterized by shorter carpels with thick peltate heads bearing two ovules on the morphologically lower surface. The cones of cycads attain in some cases (e.g.Encephalartos) a considerable size, reaching a length of more than a foot. Cases have been recorded (by Thiselton-Dyer inEncephalartosand by Wieland inZamia) in which the short carpellary cone-scales exhibit a foliaceous form. It is interesting that no monstrous cycadean cone has been described in which ovuliferous and staminate appendages are borne on the same axis: in the Bennettitales (seePalaeobotany:Mesozoic) flowers were produced bearing on the same axis both androecium and gynoecium.Fig. 7.—Zamia.Part of Ovule in longitudinal section. (After Webber.)P, Prothallus.A, Archegonia.N, Nucellus.C, Pollen-chamber.Pt, Pollen-tube.Pg, Pollen-grain.G, Generative cell (second cell of pollen-tube).Fig. 8.—Zamia.Proximal end of Pollen-tube,a,a, Spermatozoids fromGof fig. 7;Pg, pollen-grain;c, proximal cell (first cell). (After Webber.)The pollen-grains when mature consist of three cells, two small and one large cell; the latter grows into the pollen-tube, as in the Coniferales, and from one of the small cells two large ciliated spermatozoids are eventually produced. AMicrospores and megaspores.remarkable exception to this rule has recently been recorded by Caldwell, who found that inMicrocycas Calocomathe body-cells may be eight or even ten in number and the sperm-cells twice as numerous. One of the most important discoveries made during the latter part of the 19th century was that by Ikeno, a Japanese botanist, who first demonstrated the existence of motile male cells in the genusCycas. Similar spermatozoids were observed in some species ofZamiaby H. J. Webber, and more recent work enables us to assume that all cycads produce ciliated male gametes. Before following the growth of the pollen-grain after pollination, we will briefly describe the structure of a cycadean ovule. An ovule consists of a conical nucellus surrounded by a single integument. At an early stage of development a large cell makes its appearance in the central region of the nucellus; this increases in size and eventually forms three cells; the lowest of these grows vigorously and constitutes the megaspore (embryo-sac), which ultimately absorbs the greater part of the nucellus. The megaspore-nucleus divides repeatedly, and cells are produced from the peripheral region inwards, which eventually fill the spore-cavity with a homogeneous tissue (prothallus); some of the superficial cells at the micropylar end of the megaspore increase in size and divide by a tangential wall into two, an upper cell which gives rise to the short two-celled neck of the archegonium, and a lower cell which develops into a large egg-cell. Each megaspore may contain 2 to 6 archegonia. During the growth of the ovum nourishment is supplied from the contents of the cells immediately surrounding the egg-cell, as in the development of the ovum ofPinusand other conifers. Meanwhile the tissue in the apical region of the nucellus has been undergoing disorganization, which results in the formation of a pollen-chamber (fig. 7,C) immediately above the megaspore. Pollination in cycads has always been described as anemophilous, but according to recent observations by Pearson on South African species it seems probable that, at least in some cases, the pollen is conveyed to the ovules by animal agency. The pollen-grains find their way between the carpophylls, which at the time of pollination are slightly apart owing to the elongation of the internodes of the flower-axis, and pass into the pollen-chamber; the large cell of the pollen-grain grows out into a tube (Pt), which penetrates the nucellar tissue and often branches repeatedly; the pollen-grain itself, with the prothallus-cells, projects freely into the pollen-chamber (fig. 7). The nucleus of the outermost (second) small cell (fig. 7,G) divides, and one of the daughter-nuclei passes out of the cell, and may enter the lowest (first) small cell. The outermost cell, by the division of the remaining nucleus, produces two large spermatozoids (fig. 8,a,a). InMicrocycas16 sperm-cells are produced. In the course of division two bodies appear in the cytoplasm, and behave as centrosomes during the karyokinesis; they gradually become threadlike and coil round each daughter nucleus. This thread gives rise to a spiral ciliated band lying in a depression on the body of each spermatozoid; the large spermatozoids eventually escape from the pollen-tube, and are able to perform ciliary movements in the watery liquid which occurs between the thin papery remnant of nucellar tissue and the archegonial necks. Before fertilization a neck-canal cell is formed by the division of the ovum-nucleus. After the body of a spermatozoid has coalesced with the egg-nucleus the latter divides repeatedly and forms a mass of tissue which grows more vigorously in the lower part of the fertilized ovum, and extends upwards towards the apex of the ovum as a peripheral layer of parenchyma surrounding a central space. By further growth this tissue gives rise to a proembryo, which consists, at the micropylar end, of a sac; the tissue at the chalazal end grows into a long and tangled suspensor, terminating in a mass of cells, which is eventually differentiated into a radicle, plumule and two cotyledons. In the ripe seed the integument assumes the form of a fleshy envelope, succeeded internally by a hard woody shell, internal to which is a thin papery membrane—the apical portion of the nucellus—which is easily dissected out as a conical cap covering the apex of the endosperm. A thorough examination of cycadean seeds has recently been made by Miss Stopes, more particularly with a view to a comparison of their vascular supply with that in Palaeozoic gymnospermous seeds (Flora, 1904). The first leaves borne on the seedling axis are often scale-like, and these are followed by two or more larger laminae, which foreshadow the pinnae of the adult frond.Fig. 9.—Macrozamia.Diagrammatic transverse section of part of Stem. (After Worsdell.)pd, Periderm in leaf-bases.lt, Leaf-traces in cortex.ph, Phloem.x, Xylem.m, Medullary bundles.c, Cortical bundles.Fig. 10.—Ginkgo biloba.Leaves.Fig. 11.—Ginkgo adiantoides.Fossil (Eocene) leaf from the Island of Mull.The anatomical structure of the vegetative organs of recent cycads is of special interest as affording important evidence of relationship with extinct types, and with other groups of recent plants. Brongniart, whoAnatomy.was the first to investigate in detail the anatomy of a cycadean stem, recognized an agreement, as regards the secondary wood, with Dicotyledons and Gymnosperms, rather than with Monocotyledons. He drew attention also to certain structural similarities betweenCycasandGinkgo. The main anatomical features of a cycad stem may be summarized as follows: the centre is occupied by a large parenchymatous pith traversed by numerous secretory canals, and in some genera by cauline vascular bundles (e.g.EncephalartosandMacrozamia). In addition to these cauline strands (confined to the stem and not connected with the leaves), collateral bundles are often met with in the pith, which form the vascular supply of terminal flowers borne at intervals on the apex of the stem. These latter bundles may be seen in sections of old stems to pursue a more or less horizontal course, passing outwards through the main woody cylinder. This lateral course is due to the more vigorous growth of the axillary branch formed near the base of each flower, which is a terminal structure, and, except in the female flower ofCycas, puts a limit to the apical growth of the stem. The vigorous lateral branch therefore continues the line of the main axis. The pith is encircled by a cylinder of secondary wood, consisting of single or multiple radial rows of tracheids separated by broad medullary rays composed of large parenchymatous cells; the tracheids bear numerous borderedpits on the radial walls. The large medullary rays give to the wood a characteristic parenchymatous or lax appearance, which is in marked contrast to the more compact wood of a conifer. The protoxylem-elements are situated at the extreme inner edge of the secondary wood, and may occur as small groups of narrow, spirally-pitted elements scattered among the parenchyma which abuts on the main mass of wood. Short and reticulately-pitted tracheal cells, similar to tracheids, often occur in the circummedullary region of cycadean stems. In an old stem ofCycas,EncephalartosorMacrozamiathe secondary wood consists of several rather unevenly concentric zones, while in some other genera it forms a continuous mass as in conifers and normal dicotyledons. These concentric rings of secondary xylem and phloem (fig. 9) afford a characteristic cycadean feature. After the cambium has been active for some time producing secondary xylem and phloem, the latter consisting of sieve-tubes, phloem-parenchyma and frequently thick-walled fibres, a second cambium is developed in the pericycle; this produces a second vascular zone, which is in turn followed by a third cambium, and so on, until several hollow cylinders are developed. It has been recently shown that several cambium-zones may remain in a state of activity, so that the formation of a new cambium does not necessarily mark a cessation of growth in the more internal meristematic rings. It occasionally happens that groups of xylem and phloem are developed internally to some of the vascular rings; these are characterized by an inverse orientation of the tissues, the xylem being centrifugal and the phloem centripetal in its development. The broad cortical region, which contains many secretory canals, is traversed by numerous vascular bundles (fig. 9,c) some of which pursue a more or less vertical course, and by frequent anastomoses with one another form a loose reticulum of vascular strands; others are leaf-traces on their way from the stele of the stem to the leaves. Most of these cortical bundles are collateral in structure, but in some the xylem and phloem are concentrically arranged; the secondary origin of these bundles from procambium-strands was described by Mettenius in his classical paper of 1860. During the increase in thickness of a cycadean stem successive layers of cork-tissue are formed by phellogens in the persistent bases of leaves (fig. 9,pd), which increase in size to adapt themselves to the growth of the vascular zones. The leaf-traces of cycads are remarkable both on account of their course and their anatomy. In a transverse section of a stem (fig. 9) one sees some vascular bundles following a horizontal or slightly oblique course in the cortex, stretching for a longer or shorter distance in a direction concentric with the woody cylinder. From each leaf-base two main bundles spread right and left through the cortex of the stem (fig. 9,lt), and as they curve gradually towards the vascular ring they present the appearance of two rather flat ogee curves, usually spoken of as the leaf-trace girdles (fig. 9,lt). The distal ends of these girdles give off several branches, which traverse the petiole and rachis as numerous collateral bundles. The complicated girdle-like course is characteristic of the leaf-traces of most recent cycads, but in some cases,e.g.inZamia floridana, the traces are described by Wieland in his recent monograph on American fossil cycads (Carnegie Institution Publications, 1906) as possessing a more direct course similar to that in Mesozoic genera. A leaf-trace, as it passes through the cortex, has a collateral structure, the protoxylem being situated at the inner edge of the xylem; when it reaches the leaf-base the position of the spiral tracheids is gradually altered, and the endarch arrangement (protoxylem internal) gives place to a mesarch structure (protoxylem more or less central and not on the edge of the xylem strand). In a bundle examined in the basal portion of a leaf the bulk of the xylem is found to be centrifugal in position, but internally to the protoxylem there is a group of centripetal tracheids; higher up in the petiole the xylem is mainly centripetal, the centrifugal wood being represented by a small arc of tracheids external to the protoxylem and separated from it by a few parenchymatous elements. Finally, in the pinnae of the frond the centrifugal xylem may disappear, the protoxylem being now exarch in position and abutting on the phloem. Similarly in the sporophylls of some cycads the bundles are endarch near the base and mesarch near the distal end of the stamen or carpel. The vascular system of cycadean seedlings presents some features worthy of note; centripetal xylem occurs in the cotyledonary bundles associated with transfusion-tracheids. The bundles from the cotyledons pursue a direct course to the stele of the main axis, and do not assume the girdle-form characteristic of the adult plant. This is of interest from the point of view of the comparison of recent cycads with extinct species (Bennettites), in which the leaf-traces follow a much more direct course than in modern cycads. The mesarch structure of the leaf-bundles is met with in a less pronounced form in the flower peduncles of some cycads. This fact is of importance as showing that the type of vascular structure, which characterized the stems of many Palaeozoic genera, has not entirely disappeared from the stems of modern cycads; but the mesarch bundle is now confined to the leaves and peduncles. The roots of some cycadsRoots.resemble the stems in producing several cambium-rings; they possess 2 to 8 protoxylem-groups, and are characterized by a broad pericyclic zone. A common phenomenon in cycads is the production of roots which grow upwards (apogeotropic), and appear as coralline branched structures above the level of the ground; some of the cortical cells of these roots are hypertrophied, and contain numerous filaments of blue-green Algae (Nostocaceae), which live as endoparasites in the cell-cavities.Fig. 12.—Ginkgo biloba.A, Male flower;B,C, single stamens;D, female flower.Ginkgoales.—This class-designation has been recently proposed to give emphasis to the isolated position of the genusGinkgo(Salisburia) among the Gymnosperms.Ginkgo biloba, the maidenhair tree, has usually been placed by botanists in the Taxeae in the neighbourhood of the yew (Taxus), but the proposal by Eichler in 1852 to institute a special family, theSalisburieae, indicated a recognition of the existence of special characteristics which distinguish the genus from other members of the Coniferae. The discovery by the Japanese botanist Hirase of the development of ciliated spermatozoids in the pollen-tube ofGinkgo, in place of the non-motile male cells of typical conifers, served as a cogent argument in favour of separating the genus from the Coniferales and placing it in a class of its own. In 1712 Kaempfer published a drawing of a Japanese tree, which he described under the nameGinkgo; this term was adopted in 1771 by Linnaeus, who spoke of Kaempfer’s plant asGinkgo biloba. In 1797 Smith proposed to use the nameSalisburia adiantifoliain preference to the “uncouth” genusGinkgoand “incorrect” specific termbiloba. Both names are still in common use. On account of the resemblance of the leaves to those of some species ofAdiantum, the appellation maidenhair tree has long been given toGinkgo biloba.Ginkgois of special interest on account of its isolated position among existing plants, its restricted geographical distribution, and its great antiquity (seePalaeobotany:Mesozoic). This solitary survivor of an ancient stock is almost extinct, but a few old and presumably wild trees are recorded by travellers in parts of China.Ginkgois common as a sacred tree in the gardens of temples in the Far East, and often cultivated in North America and Europe.Ginkgo biloba, which may reach a height of over 30 metres, forms a tree of pyramidal shape with a smooth grey bark. The leaves (figs. 10 and 11) have a long, slender petiole terminating in a fan-shaped lamina, which may be entire, divided by a median incision into two wedge-shaped lobes, or subdivided into several narrow segments. The venation is like that of many ferns,e.g.Adiantum; the lowest vein in each half of the lamina follows a course parallel to the edge, and gives off numerous branches, which fork repeatedly as they spread in a palmate manner towards the leaf margin. The foliage-leaves occur either scattered on long shoots of unlimited growth, or at the apex of short shoots (spurs), which may eventually elongate into long shoots.Fig. 13.—Ginkgo.Apex of Ovule, and Pollen-grain. (After Hirase.)p, Pollen-tube (proximal end).c, Pollen-chamber.e, Upward prolongation of megaspore.a, Archegonia.Pg, Pollen-grain.Ex, Exine.The flowers are dioecious. The male flowers (fig. 12), borne in the axil of scale-leaves, consist of a stalked central axis bearing loosely disposed stamens; each stamen consists of a slender filament terminating in a small apical scale, which bearsFlowers.usually two, but not infrequently three or four pollen-sacs (fig. 12,C). The axis of the flower is a shoot bearing leaves in the form of stamens. A mature pollen-grain contains a prothallus of 3 to 5 cells (Fig. 13, Pg); the exine extends over two-thirds of the circumference, leaving a thin portion of the wall, which on collapsing produces a longitudinal groove similar to the median depression on the pollen-grain of a cycad. The ordinary type of female flower has the form of a long, naked peduncle bearing a single ovule on either side of the apex (fig. 12), the base of each being enclosed by a small, collar-like rim, the nature of which has been variously interpreted. A young ovule consists of a conical nucellus surrounded by a single integument terminating as a two-lipped micropyle. A large pollen-chamber occupies the apex of the nucellus; immediately below this, two or more archegonia (fig. 13,a) are developed in the upper region of the megaspore, each consisting of a large egg-cell surmounted by two neck-cells and a canal-cell which is cut off shortly before fertilization. After the entrance of the pollen-grain the pollen-chamber becomes roofed over by a blunt protuberance of nucellar tissue. The megaspore (embryo-sac) continues to grow after pollination until the greater part of the nucellus is gradually destroyed; it also gives rise to a vertical outgrowth, which projects from the apex of the megaspore as a short, thick column (fig. 13,e) supporting the remains of the nucellar tissue which forms the roof of the pollen-chamber (fig. 13,c). Surrounding the pitted wall of the ovum there is a definite layer of large cells, no doubt representing a tapetum, which, as in cycads and conifers, plays an important part in nourishing the growing egg-cell. The endosperm detached from a largeGinkgoovule after fertilization bears a close resemblance to that of a cycad; the apex is occupied by a depression, on the floor of which two small holes mark the position of the archegonia, and the outgrowth from the megaspore apex projects from the centre as a short peg. After pollination the pollen-tube grows into the nucellar tissue, as in cycads, and the pollen-grain itself (fig. 13,Pg) hangs down into the pollen-chamber; two large spirally ciliated spermatozoids are produced, their manner of development agreeing very closely with that of the corresponding cells inCycasandZamia. After fertilization the ovum-nucleus divides and cell-formation proceeds rapidly, especially in the lower part of the ovum, in which the cotyledon and axis of the embryo are differentiated; the long, tangled suspensor of the cycadean embryo is not found inGinkgo. It is often stated that fertilization occurs after the ovules have fallen, but it has been demonstrated by Hirase that this occurs while the ovules are still attached to the tree. The ripe seed, which grows as large as a rather small plum, is enclosed by a thick, fleshy envelope covering a hard woody shell with two or rarely three longitudinal keels. A papery remnant of nucellus lines the inner face of the woody shell, and, as in cycadean seeds, the apical portion is readily separated as a cap covering the summit of the endosperm.Fig. 14.—Ginkgo. Abnormal female Flowers.A, Peduncle;b, scaly bud;B, leaf bearing marginal ovule. (After Fujii.)The morphology of the female flowers has been variously interpreted by botanists; the peduncle bearing the ovules has been described as homologous with the petiole of a foliage-leaf and as a shoot-structure, the collar-like envelope at the base of the ovules being referred to as a second integument or arillus, or as the representative of a carpel. The evidence afforded by normal and abnormal flowers appears to be in favour of the following interpretation: The peduncle is a shoot bearing two or more carpels. Each ovule is enclosed at the base by an envelope or collar homologous with the lamina of a leaf; the fleshy and hard coats of the nucellus constitute a single integument. The stalk of an ovule, considerably reduced in normal flowers and much larger in some abnormal flowers, is homologous with a leaf-stalk, with which it agrees in the structure and number of vascular bundles. The facts on which this description is based are derived partly from anatomical evidence, and in part from an account given by a Japanese botanist, Fujii, of several abnormal female flowers; in some cases the collar at the base of an ovule, often described as an arillus, is found to pass gradually into the lamina of a leaf bearing marginal ovules (fig. 14,B). The occurrence of more than two ovules on one peduncle is by no means rare; a particularly striking example is described by Fujii, in which an unusually thick peduncle bearing several stalked ovules terminates in a scaly bud (fig. 14,A,b). The frequent occurrence of more than two pollen-sacs and the equally common occurrence of additional ovules have been regarded by some authors as evidence in favour of the view that ancestral types normally possessed a greater number of these organs than are usually found in the recent species. ThisAnatomy.view receives support from fossil evidence. Close to the apex of a shoot the vascular bundles of a leaf make their appearance as double strands, and the leaf-traces in the upper part of a shoot have the form of distinct bundles, which in the older part of the shoot form a continuous ring. Each double leaf-trace passes through four internodes before becoming a part of the stele; the double nature of the trace is a characteristic feature. Secretory sacs occur abundantly in the leaf-lamina, where they appear as short lines between the veins; they are abundant also in the cortex and pith of the shoot, in the fleshy integument of the ovule, and elsewhere. The secondary wood of the shoot and root conforms in the main to the coniferous type; in the short shoots the greater breadth of the medullary rays in the more internal part of the xylem recalls the cycadean type. The secondary phloem contains numerous thick-walled fibres, parenchymatous cells, and large sieve-tubes with plates on the radial walls; swollen parenchymatous cells containing crystals are commonly met with in the cortex, pith and medullary-ray tissues. The wood consists of tracheids, with circular bordered pits on their radial walls, and in the late summer wood pits are unusually abundant on the tangential walls. A point of anatomical interest is the occurrence in the vascular bundles of the cotyledons, scale-leaves, and elsewhere of a few centripetally developed tracheids, which give to the xylem-strands a mesarch structure such as characterizes the foliar bundles of cycads. The root is diarch in structure, but additional protoxylem-strands may be present at the base of the main root; the pericycle consists of several layers of cells.This is not the place to discuss in detail the past history ofGinkgo(seePalaeobotany:Mesozoic). Among Palaeozoic genera there are some which bear a close resemblance to the recent type in the form of the leaves; and petrified Palaeozoic seeds,Geological history.almost identical with those of the maidenhair tree, have been described from French and English localities. During the Triassic and Jurassic periods the genusBaiera—no doubt a representative of the Ginkgoales—was widely spread throughout Europe and in other regions;Ginkgoitself occurs abundantly in Mesozoic and Tertiary rocks, and was a common plant in the Arctic regions as elsewhere during the Jurassic and Lower Cretaceous periods. Some unusually perfectGinkgoleaves have been found in the Eocene leaf-beds between the lava-flows exposed in the cliffs of Mull (fig. 11). From an evolutionary point of view, it is of interest to note the occurrence of filicinean and cycadean characters in the maidenhair tree. The leaves at once invite a comparison with ferns; the numerous long hairs which form a delicate woolly covering on young leaves recall the hairs of certain ferns, but agree more closely with the long filamentous hairs of recent cycads. The spermatozoids constitute the most striking link with both cycads and ferns. The structure of the seed, the presence of two neck-cells in the archegonia, the late development of the embryo, the partially-fused cotyledons and certain anatomical characters, are features common toGinkgoand the cycads. The maidenhair tree is one of the most interesting survivals from the past; it represents a type which, in the Palaeozoic era, may have been merged into the extinct class Cordaitales. Through the succeeding ages the Ginkgoales were represented by numerous forms, which gradually became more restricted in their distribution and fewer in number during the Cretaceous and Tertiary periods, terminating at the present day in one solitary survivor.Coniferales.—Trees and shrubs characterized by a copious branching of the stem and frequently by a regular pyramidal form. Leaves simple, small, linear or short and scale-like, usually persisting for more than one year. Flowers monoecious or dioecious, unisexual, without a perianth, often in the form of cones, but never terminal on the main stem.The plants usually included in the Coniferae constitute a less homogeneous class than the Cycadaceae. Some authors use the term Coniferae in a restricted sense as including those genera which have the female flowers in the form of cones,External features.the other genera, characterized by flowers of a different type, being placed in the Taxaceae, and often spoken of as Taxads.In order to avoid confusion in the use of the term Coniferae, we may adopt as a class-designation the name Coniferales, including both the Coniferae—using the term in a restricted sense—and the Taxaceae. The most striking characteristic of the majority of the Coniferales is the regular manner of the monopodial branching and the pyramidal shape.Araucaria imbricata, the Monkey-puzzle tree,A. excelsa, the Norfolk Island pine, many pines and firs, cedars and other genera illustrate the pyramidal form. The mammoth redwood tree of California,Sequoia(Wellingtonia)gigantea, which represents the tallest Gymnosperm, is a good example of the regular tapering main stem and narrow pyramidal form. The cypresses afford instances of tall and narrow trees similar in habit to Lombardy poplars. The common cypress (Cupressus sempervirens), as found wild in the mountains of Crete and Cyprus, is characterized by long and spreading branches, which give it a cedar-like habit. A pendulous or weeping habit is assumed by some conifers,e.g.Picea excelsavar.virgatarepresents a form in which the main branches attain a considerable horizontal extension, and trail themselves like snakes along the ground. Certain species ofPinus, the yews (Taxus) and some other genera grow as bushes, which in place of a main mast-like stem possess several repeatedly-branched leading shoots. The unfavourable conditions in Arctic regions have produced a dwarf form, in which the main shoots grow close to the ground. Artificially induced dwarfed plants ofPinus,Cupressus,Sciadopitys(umbrella pine) and other genera are commonly cultivated by the Japanese. The dying off of older branches and the vigorous growth of shoots nearer the apex of the stem produce a form of tree illustrated by the stone pine of the Mediterranean region (Pinus Pinea), which Turner has rendered familiar in his “Childe Harold’s Pilgrimage” and other pictures of Italian scenery. Conifers are not infrequently seen in which a lateral branch has bent sharply upwards to take the place of the injured main trunk. An upward tendency of all the main lateral branches, known as fastigiation, is common in some species, producing well-marked varieties,e.g.Cephalotaxus pedunculatavar.fastigiata; this fastigiate habit may arise as a sport on a tree with spreading branches. Another departure from the normal is that in which the juvenile or seedling form of shoot persists in the adult tree; the numerous coniferous plants known as species ofRetinosporaare examples of this. The nameRetinospora, therefore, does not stand for a true genus, but denotes persistent young forms ofJuniperus,Thuja,Cupressus, &c., in which the small scaly leaves of ordinary species are replaced by the slender, needle-like leaves, which stand out more or less at right angles from the branches. The flat branchlets ofCupressus,Thuja(arbor vitae),Thujopsis dolabrata(Japanese arbor vitae) are characteristic of certain types of conifers; in some cases the horizontal extension of the branches induces a dorsiventral structure. A characteristic feature of the genusAgathis(Dammara) the Kauri pine of New Zealand, is the deciduous habit of the branches; these become detached from the main trunk leaving a well-defined absciss-surface, which appears as a depressed circular scar on the stem. A new genus of conifers,Taiwania, has recently been described from the island of Formosa; it is said to agree in habit with the JapaneseCryptomeria, but the cones appear to have a structure which distinguishes them from those of any other genus.With a few exceptions conifers are evergreen, and retain the leaves for several years (10 years inAraucaria imbricata, 8 to 10 inPicea excelsa, 5 inTaxus baccata; inPinusthe needles usually fall in October of their third year). The larch (Larix)Leaves.sheds its leaves in the autumn, in the Chinese larch (Pseudolarix Kaempferi) the leaves turn a bright yellow colour before falling. In the swamp cypress (Taxodium distichum) the tree assumes a rich brown colour in the autumn, and sheds its leaves together with the branchlets which bear them; deciduous branches occur also in some other species,e.g.Sequoia sempervirens(redwood),Thuja occidentalis, &c. The leaves of conifers are characterized by their small size,e.g.the needle-form represented byPinus,Cedrus,Larix, &c., the linear flat or angular leaves, appressed to the branches, ofThuja,Cupressus,Libocedrus, &c. The flat and comparatively broad leaves ofAraucaria imbricata,A. Bidwillii, and some species of the southern genusPodocarpusare traversed by several parallel veins, as are also the still larger leaves ofAgathis, which may reach a length of several inches. In addition to the foliage-leaves several genera also possess scale-leaves of various kinds, represented by bud-scales inPinus,Picea, &c., which frequently persist for a time at the base of a young shoot which has pushed its way through the yielding cap of protecting scales, while in some conifers the bud-scales adhere together, and after being torn near the base are carried up by the growing axis as a thin brown cap. The cypresses, araucarias and some other genera have no true bud-scales; in some species,e.g.Araucaria Bidwillii, the occurrence of small foliage-leaves, which have functioned as bud-scales, at intervals on the shoots affords a measure of seasonal growth. The occurrence of long and short shoots is a characteristic feature of many conifers. InPinusthe needles occur in pairs, or in clusters of 3 or 5 at the apex of a small and inconspicuous short shoot of limited growth (spur), which is enclosed at its base by a few scale-leaves, and borne on a branch of unlimited growth in the axil of a scale-leaf. In the CalifornianPinus monophyllaeach spur bears usually one needle, but two are not uncommon; it would seem that rudiments of two needles are always produced, but, as a rule, only one develops into a needle. InSciadopityssimilar spurs occur, each bearing a single needle, which in its grooved surface and in the possession of a double vascular bundle bears traces of an origin from two needle-leaves. A peculiarity of these leaves is the inverse orientation of the vascular tissue; each of the two veins has its phloem next the upper and the xylem towards the lower surface of the leaf; this unusual position of the xylem and phloem may be explained by regarding the needle ofSciadopitysas being composed of a pair of leaves borne on a short axillary shoot and fused by their margins (fig. 15, A). Long and short shoots occur also inCedrusandLarix, but in these genera the spurs are longer and stouter, and are not shed with the leaves; this kind of short shoot, by accelerated apical growth, often passes into the condition of a long shoot on which the leaves are scattered and separated by comparatively long internodes, instead of being crowded into tufts such as are borne on the ends of the spurs. In the genusPhyllocladus(New Zealand, &c.) there are no green foliage-leaves, but in their place flattened branches (phylloclades) borne in the axils of small scale-leaves. The cotyledons are often two in number, but sometimes (e.g.Pinus) as many as fifteen; these leaves are usually succeeded by foliage-leaves in the form of delicate spreading needles, and these primordial leaves are followed, sooner or later, by the adult type of leaf, except in Retinosporas, which retain the juvenile foliage. In addition to the first foliage-leaves and the adult type of leaf, there are often produced leaves which are intermediate both in shape and structure between the seedling and adult foliage. Dimorphism or heterophylly is fairly common. One of the best known examples is the Chinese juniper (Juniperus chinensis), in which branches with spinous leaves, longer and more spreading than the ordinary adult leaf, are often found associated with the normal type of branch. In some cases,e.g.Sequoia sempervirens, the fertile branches bear leaves which are less spreading than those on the vegetative shoots. Certain species of the southern hemisphere genusDacrydiumafford particularly striking instances of heterophylly,e.g.D. Kirkiiof New Zealand, in which some branches bear small and appressed leaves, while in others the leaves are much longer and more spreading. A well-known fossil conifer from Triassic strata—Voltzia heterophylla—also illustrates a marked dissimilarity in the leaves of the same shoot. The variation in leaf-form and the tendency of leaves to arrange themselves in various ways on different branches of the same plant are features which it is important to bear in mind in the identification of fossil conifers. In this connexion we may note the striking resemblance between some of the New Zealand Alpine Veronicas,e.g.Veronica Hectori,V. cupressoides, &c. (alsoPolycladus cupressinus, a Composite), and some of the cypresses and other conifers with small appressed leaves. The long linear leaves of some species ofPodocarpus, in which the lamina is traversed by a single vein, recall the pinnae of Cycas; the branches of some Dacrydiums and other forms closely resemble those of lycopods; these superficial resemblances, both between different genera of conifers and between conifers and other plants, coupled with the usual occurrence of fossil coniferous twigs without cones attached to them, render the determination of extinct types a very unsatisfactory and frequently an impossible task.A typical male flower consists of a central axis bearing numerous spirally-arranged sporophylls (stamens), each of which consists of a slender stalk (filament) terminating distally in a more or less prominent knob or triangular scale, and bearingFlowers.two or more pollen-sacs (microsporangia) on its lower surface. The pollen-grains of some genera (e.g.Pinus) are furnished with bladder-like extensions of the outer wall, which serve as aids to wind-dispersal. The stamens ofAraucariaandAgathisare peculiar in bearing several long, and narrow free pollen-sacs; these may be compared with the sporangiophores of the horsetails (Equisetum); inTaxus(yew) the filament is attached to the centre of a large circular distal expansion, which bears several pollen-sacs on its under surface. In the conifers proper the female reproductive organs have the form of cones, which may be styled flowers or inflorescences according to different interpretations of their morphology. In the Taxaceae the flowers have a simpler structure. The female flowers of theAbietineaemay be taken as representing a common type. A pine cone reaches maturity in two years; a single year suffices for the full development inLarixand several other genera. The axis of the cone bears numerous spirally disposed flat scales (cone-scales), each of which, if examined in a young cone, is found to be double, and to consist of a lower and an upper portion. The latter is a thin flat scale bearing a median ridge or keel (e.g.Abies), on each side of which is situated an inverted ovule, consisting of a nucellus surrounded by a single integument. As the cone grows in size and becomes woody the lower half of the cone-scale, which we may call the carpellary scale, may remain small, and is so far outgrown by the upper half (seminiferous scale) that it is hardly recognizable in the mature cone. In many species ofAbies(e.g.Abies pectinata, &c.) the ripe cone differs from those ofPinus,PiceaandCedrusin the large size of the carpellary scales, which project as conspicuous thin appendages beyond the distal margins of the broader and more woody seminiferous scales; the long carpellary scale is a prominent feature also in the cone of the Douglas pine (Pseudotsuga Douglasii). The female flowers (cones) vary considerably in size; the largest are the more or less spherical cones ofAraucaria—a single cone ofA. imbricatamay produce as many as 300 seeds, one seed to each fertile cone-scale—and the long pendentcones, 1 to 2 ft. in length, of the sugar pine of California (Pinus Lambertiana) and other species. Smaller cones, less than an inch long, occur in the larch,Athrotaxis(Tasmania),Fitzroya(Patagonia and Tasmania), &c. In theTaxodieaeandAraucarieaethe cones are similar in appearance to those of theAbietineae, but they differ in the fact that the scales appear to be single, even in the young condition; each cone-scale in a genus of theTaxodiinae(Sequoia, &c.) bears several seeds, while in theAraucariinae(AraucariaandAgathis) each scale has one seed. TheCupressineaehave cones composed of a few scales arranged in alternate whorls; each scale bears two or more seeds, and shows no external sign of being composed of two distinct portions. In the junipers the scales become fleshy as the seeds ripen, and the individual scales fuse together in the form of a berry. The female flowers of the Taxaceae assume another form; inMicrocachrys(Tasmania) the reproductive structures are spirally disposed, and form small globular cones made up of red fleshy scales, to each of which is attached a single ovule enclosed by an integument and partially invested by an arillus; inDacrydiumthe carpellary leaves are very similar to the foliage leaves—each bears one ovule with two integuments, the outer of which constitutes an arillus. Finally in the yew, as a type of the family Taxeae, the ovules occur singly at the apex of a lateral branch, enclosed when ripe by a conspicuous red or yellow fleshy arillus, which serves as an attraction to animals, and thus aids in the dispersal of the seeds.(C and D after Worsdell.)Fig. 15.—Diagrammatic treatment of:A, Double needle ofSciadopitys(a,a, leaves;b, shoot;Br, bract).B, seminiferous scale as leaf of axillary shoot (b, shoot;Sc, seminiferous scale;Br, bract).C, seminiferous scale as fused pair of leaves (l1,l2,l3, first, second and third leaves;b, shoot;Br, bract).D, cone-scale ofAraucaria(n, nucellus;i, integument;x, xylem).Fig. 16.—Abnormal Cone ofPinus rigida. (After Masters.)It is important to draw attention to some structural features exhibited by certain cone-scales, in which there is no external sign indicative of the presence of a carpellary and a seminiferous scale. InAraucaria Cookiiand some allied species eachMorphology of female flower.scale has a small pointed projection from its upper face near the distal end; the scales ofCunninghamia(China) are characterized by a somewhat ragged membranous projection extending across the upper face between the seeds and the distal end of the scale; in the scales ofAthrotaxis(Tasmania) a prominent rounded ridge occupies a corresponding position. These projections and ridges may be homologous with the seminiferous scale of the pines, firs, cedars, &c. The simplest interpretation of the cone of theAbietineaeis that which regards it as a flower consisting of an axis bearing several open carpels, which in the adult cone may be large and prominent or very small, the scale bearing the ovules being regarded as a placental outgrowth from the flat and open carpel. InAraucariathe cone-scale is regarded as consisting of a flat carpel, of which the placenta has not grown out into the scale-like structure. The seminiferous scale ofPinus, &c., is also spoken of sometimes as a ligular outgrowth from the carpellary leaf. Robert Brown was the first to give a clear description of the morphology of the Abietineous cone in which carpels bear naked ovules; he recognized gymnospermy as an important distinguishing feature in conifers as well as in cycads. Another view is to regard the cone as an inflorescence, each carpellary scale being a bract bearing in its axil a shoot the axis of which has not been developed; the seminiferous scale is believed to represent either a single leaf or a fused pair of leaves belonging to the partially suppressed axillary shoot. In 1869 van Tieghem laid stress on anatomical evidence as a key to the morphology of the cone-scales; he drew attention to the fact that the collateral vascular bundles of the seminiferous scale are inversely orientated as compared with those of the carpellary scale; in the latter the xylem of each bundle is next the upper surface, while in the seminiferous scale the phloem occupies that position. The conclusion drawn from this was that the seminiferous scale (fig. 15, B,Sc) is the first and only leaf of an axillary shoot (b) borne on that side of the shoot, the axis of which is suppressed, opposite the subtending bract (fig. 15, A, B, C,Br). Another view is to apply to the seminiferous scale an explanation similar to that suggested by von Mohl in the case of the double needle ofSciadopitys, and to consider the seed-bearing scale as being made up of a pair of leaves (fig. 15, A,a,a) of an axillary shoot (b) fused into one by their posterior margins (fig. 15, A). The latter view receives support from abnormal cones in which carpellary scales subtend axillary shoots, of which the first two leaves (fig. 15, C,l1,l1) are often harder and browner than the others; forms have been described transitional between axillary shoots, in which the leaves are separate, and others in which two of the leaves are more or less completely fused. In a young cone the seminiferous scale appears as a hump of tissue at the base or in the axil of the carpellary scale, but Celakovský, a strong supporter of the axillary-bud theory, attaches little or no importance to this kind of evidence, regarding the present manner of development as being merely an example of a short cut adopted in the course of evolution, and replacing the original production of a branch in the axil of each carpellary scale. Eichler, one of the chief supporters of the simpler view, does not recognize in the inverse orientation of the vascular bundles an argument in support of the axillary-bud theory, but points out that the seminiferous scale, being an outgrowth from the surface of the carpellary scale, would, like outgrowths from an ordinary leaf, naturally have its bundles inversely orientated. In such cone-scales as show little or no external indication of being double in origin,e.g.Araucaria(fig. 15, D) Sequoia, &c., there are always two sets of bundles; the upper set, having the phloem uppermost, as in the seminiferous scale ofAbiesorPinus, are regarded as belonging to the outgrowth from the carpellary scale and specially developed to supply the ovules. Monstrous cones are fairly common; these in some instances lend support to the axillary-bud theory, and it has been said that this theory owes its existence to evidence furnished by abnormal cones. It is difficult to estimate the value of abnormalities as evidence bearing on morphological interpretation; the chief danger lies perhaps in attaching undue weight to them, but there is also a risk of minimizing their importance. Monstrosities at least demonstrate possible lines of development, but when the abnormal forms of growth in various directions are fairly evenly balanced, trustworthy deductions become difficult. The occurrence of buds in the axils of carpellary scales may, however, simply mean that buds, which are usually undeveloped in the axils of sporophylls, occasionally afford evidence of their existence. Some monstrous cones lend no support to the axillary-bud theory. InLarixthe axis of the cone often continues its growth; similarly inCephalotaxusthe cones are often proliferous. (In rare cases the proliferated portion produces male flowers in the leaf-axils.) InLarixthe carpellary scale may become leafy, and the seminiferous scale may disappear. Androgynous cones may be produced, as in the cone ofPinus rigida(fig. 16), in which the lower part bears stamens and the upper portion carpellary and seminiferous scales. An interesting case has been figured by Masters, in which scales of a cone ofCupressus Lawsonianabear ovules on the upper surface and stamens on the lower face. One argument that has been adduced in support of the axillary bud theory is derived from the Palaeozoic typeCordaites, in which each ovule occurs on an axis borne in the axil of a bract. The whole question is still unsolved, and perhaps insoluble. It may be that the interpretation of the female cone of theAbietineaeas an inflorescence, which finds favour with many botanists, cannot be applied to the cones ofAgathisandAraucaria. Without expressing any decided opinion as to the morphology of the double cone-scale of theAbietineae, preference may be felt in favour of regarding the cone-scale of theAraucarieaeas a simple carpellary leaf bearing a single ovule. A discussion of this question may be found in a paper on theAraucarieaeby Seward and Ford, published in the Transactions of the Royal Society of London (1906).Cordaitesis an extinct type which in certain respects resemblesGinkgo, cycads and theAraucarieae, but its agreement with true conifers is probably too remote to justify our attributing much weight to the bearing of the morphology of its female flowers on the interpretation of that of the Coniferae. The greater simplicity of the Eichler theory may prejudice us in its favour; but, on the other hand, the arguments advanced in favour of the axillary-bud theories are perhaps not sufficiently cogent to lead us to accept an explanation based chiefly on the uncertain evidence of monstrosities.A pollen-grain when first formed from its mother-cell consists of a single cell; in this condition it may be carried to the nucellus of the ovule (e.g.Taxus,Cupressus, &c.), or more usually (Pinus,Larix, &c.) it reaches maturity before the dehiscenceMicro-spores and megaspores.of the microsporangium. The nucleus of the microspore divides and gives rise to a small cell within the large cell, a second small cell is then produced; this is the structure of the ripe pollen-grain in some conifers (Taxus, &c.). The large cell grows out as a pollen-tube; the second of the two small cells (body-cell) wanders into the tube, followed by the nucleus of the first small cell (stalk-cell). InTaxusthe body-cell eventually divides into two, in which the products of division are of unequal size, the larger constituting the male generative cell, which fuses with the nucleus of the egg-cell. InJuniperusthe products of division of thebody-cell are equal, and both function as male generative cells. In theAbietineaecell-formation in the pollen-grain is carried farther. Three small cells occur inside the cavity of the microspore; two of them collapse and the third divides into two, forming a stalk-cell and a larger body-cell. The latter ultimately divides in the apex of the pollen-tube into two non-motile generative cells. Evidence has lately been adduced of the existence of numerous nuclei in the pollen-tubes of theAraucarieae, and it seems probable that in this as in several other respects this family is distinguished from other members of the Coniferales. The precise method of fertilization in the Scots Pine was followed by V. H. Blackman, who also succeeded in showing that the nuclei of the sporophyte generation contain twice as many chromosomes as the nuclei of the gametophyte. Other observers have in recent years demonstrated a similar relation in other genera between the number of chromosomes in the nuclei of the two generations. The ovule is usually surrounded by one integument, which projects beyond the tip of the nucellus as a wide-open lobed funnel, which at the time of pollination folds inwards, and so assists in bringing the pollen-grains on to the nucellus. In some conifers (e.g.Taxus,Cephalotaxus,Dacrydium, &c.) the ordinary integument is partially enclosed by an arillus or second integument. It is held by some botanists (Celakovský) that the seminiferous scale of theAbietineaeis homologous with the arillus or second integument of the Taxaceae, but this view is too strained to gain general acceptance. InAraucariaandSaxegothaeathe nucellus itself projects beyond the open micropyle and receives the pollen-grains direct. During the growth of the cell which forms the megaspore the greater part of the nucellus is absorbed, except the apical portion, which persists as a cone above the megaspore; the partial disorganization of some of the cells in the centre of the nucellar cone forms an irregular cavity, which may be compared with the larger pollen-chamber ofGinkgoand the cycads. In each ovule one megaspore comes to maturity, but, exceptionally, two may be present (e.g.Pinus sylvestris). It has been shown by Lawson that inSequoia sempervirens(Annals of Botany, 1904) and by other workers in the genera that several megaspores may attain a fairly large size in one prothallus. The megaspore becomes filled with tissue (prothallus), and from some of the superficial cells archegonia are produced, usually three to five in number, but in rare cases ten to twenty or even sixty may be present. In the genusSequoiathere may be as many as sixty archegonia (Arnoldi and Lawson) in one megaspore; these occur either separately or in some parts of the prothallus they may form groups as in theCupressineae; they are scattered through the prothallus instead of being confined to the apical region as in the majority of conifers. Similarly in theAraucarieaeand inWiddringtoniathe archegonia are numerous and scattered and often sunk in the prothallus tissue. InLibocedrus decurrens(Cupressineae) Lawson describes the archegonia as varying in number from 6 to 24 (Annals of Botanyxxi., 1907). An archegonium consists of a large oval egg-cell surmounted by a short neck composed of one or more tiers of cells, six to eight cells in each tier. Before fertilization the nucleus of the egg-cell divides and cuts off a ventral canal-cell; this cell may represent a second egg-cell. The egg-cells of the archegonia may be in lateral contact (e.g.Cupressineae) or separated from one another by a few cells of the prothallus, each ovum being immediately surrounded by a layer of cells distinguished by their granular contents and large nuclei. During the development of the egg-cell, food material is transferred from these cells through the pitted wall of the ovum. The tissue at the apex of the megaspore grows slightly above the level of the archegonia, so that the latter come to lie in a shallow depression. In the process of fertilization the two male generative nuclei, accompanied by the pollen-tube nucleus and that of the stalk-cell, pass through an open pit at the apex of the pollen-tube into the protoplasm of the ovum. After fertilization the nucleus of the egg divides, the first stages of karyokinesis being apparent even before complete fusion of the male and female nuclei has occurred. The result of this is the production of four nuclei, which eventually take up a position at the bottom of the ovum and become separated from one another by vertical cell-walls; these nuclei divide again, and finally three tiers of cells are produced, four in each tier. In theAbietineaethe cells of the middle tier elongate and push the lowest tier deeper into the endosperm; the cells of the bottom tier may remain in lateral contact and produce together one embryo, or they may separate (Pinus,Juniperus, &c.) and form four potential embryos. The ripe albuminous seed contains a single embryo with two or more cotyledons. The seeds of many conifers are provided with large thin wings, consisting in some genera (e.g.Pinus) of the upper cell-layers of the seminiferous scale, which have become detached and, in some cases, adhere loosely to the seed as a thin membrane; the loose attachment may be of use to the seeds when they are blown against the branches of trees, in enabling them to fall away from the wing and drop to the ground. The seeds of some genera depend on animals for dispersal, the carpellary scale (Microcachrys) or the outer integument being brightly coloured and attractive. In someAbietineae(e.g.PinusandPicea)—in which the cone-scales persist for some time after the seeds are ripe—the cones hang down and so facilitate the fall of the seeds; inCedrus,AraucariaandAbiesthe scales become detached and fall with the seeds, leaving the bare vertical axis of the cone on the tree. In all cases, except some species ofAraucaria(sect.Colymbea) the germination is epigean. The seedling plants of some Conifers (e.g.Araucaria imbricata) are characterized by a carrot-shaped hypocotyl, which doubtless serves as a food-reservoir.The roots of many conifers possess a narrow band of primary xylem-tracheids with a group of narrow spiral protoxylem-elements at each end (diarch). A striking feature in the roots of several genera, excluding theAbietineae, is the occurrenceAnatomy.of thick and somewhat irregular bands of thickening on the cell-walls of the cortical layer next to the endodermis. These bands, which may serve to strengthen the central cylinder, have been compared with the netting surrounding the delicate wall of an inflated balloon. It is not always easy to distinguish a root from a stem; in some cases (e.g.Sequoia) the primary tetrarch structure is easily identified in the centre of an old root, but in other cases the primary elements are very difficult to recognize. The sudden termination of the secondary tracheids against the pith-cells may afford evidence of root-structure as distinct from stem-structure, in which the radial rows of secondary tracheids pass into the irregularly-arranged primary elements next the pith. The annual rings in a root are often less clearly marked than in the stem, and the xylem-elements are frequently larger and thinner. The primary vascular bundles in a young conifer stem are collateral, and, like those of a Dicotyledon, they are arranged in a circle round a central pith and enclosed by a common endodermis. It is in the nature of the secondary xylem that the Coniferales are most readily distinguished from the Dicotyledons and Cycadaceae; the wood is homogeneous in structure, consisting almost entirely of tracheids with circular or polygonal bordered pits on the radial walls, more particularly in the late summer wood. In many genera xylem-parenchyma is present, but never in great abundance. A few Dicotyledons,e.g.Drimys(Magnoliaceae) closely resemble conifers in the homogeneous character of the wood, but in most cases the presence of large spring vessels, wood-fibres and abundant parenchyma affords an obvious distinguishing feature.
Gymnospermae.—Trees or shrubs; leaves vary considerably in size and form. Flowers unisexual, except in a few cases (Gnetales) without a perianth. Monoecious or dioecious. Ovules naked, rarely without carpellary leaves, usually borne on carpophylls, which assume various forms. The single megaspore enclosed in the nucellus is filled with tissue (prothallus) before fertilization, and contains two or more archegonia, consisting usually of a large egg-cell and a small neck, rarely of an egg-cell only and no neck (GnetumandWelwitschia). Microspore spherical or oval, with or without a bladder-like extension of the exine, containing a prothallus of two or more cells, one of which produces two non-motile or motile male cells. Cotyledons two or several. Secondary xylem and phloem produced by a single cambium, or by successive cambial zones; no true vessels (except in the Gnetales) in the wood, and no companion-cells in the phloem.
There is no doubt that the result of recent research and of work now in progress will be to modify considerably the grouping of the conifers. The familyAraucarieae, represented byAraucariaandAgathis, should perhaps be separated as a special class and a rearrangement of other genera more in accord with a natural system of classification will soon be possible; but for the present its twofold subdivision may be retained.
Cycadophyta.—A.Cycadales.—Stems tuberous or columnar, not infrequently branched, rarely epiphytic (Peruvian species ofZamia); fronds pinnate, bi-pinnate in the Australian genusBowenia. Dioecious; flowers in the form of cones, except the female flowers ofCycas, which consist of a rosette of leaf-like carpels at the apex of the stem. Seeds albuminous, with one integument; the single embryo, usually bearing two partially fused cotyledons, is attached to a long tangled suspensor. Stems and roots increase in diameter by secondary thickening, the secondary wood being produced by one cambium or developed from successive cambium-rings.
The cycads constitute a homogeneous group of a few living members confined to tropical and sub-tropical regions. As a fairly typical and well-known example of the Cycadaceae, a species of the genusCycas(e.g.C. circinalis,C. revoluta, &c.) is briefly described. The stout columnar stem may reach a height of 20 metres, and a diameter of half a metre; it remains either unbranched or divides near the summit into several short and thick branches, each branch terminating in a crown of long pinnate leaves. The surface of the stem is covered with rhomboidal areas, which represent the persistent bases of foliage- and scale-leaves. In some species ofCycasthere is a well-defined alternation of transverse zones on the stem, consisting of larger areas representing foliage-leaf bases, and similar but smaller areas formed by the bases of scale-leaves (FandS, fig. 1). The scale-leaves clothing the terminal bud are linear-lanceolate in form, and of a brown or yellow colour; they are pushed aside as the stem-axis elongates and becomes shrivelled, finally falling off, leaving projecting bases which are eventually cut off at a still lower level. Similarly, the dead fronds fall off, leaving a ragged petiole, which is afterwards separated from the stem by an absciss-layer a short distance above the base. In some species ofCycasthe leaf-bases do not persist as a permanent covering to the stem, but the surfaceis covered with a wrinkled bark, as inCycas siamensis, which has a stem of unusual form (fig. 2). Small tuberous shoots, comparable on a large scale with the bulbils ofLycopodium Selago, are occasionally produced in the axils of some of the persistent leaf-bases; these are characteristic of sickly plants, and serve as a means of vegetative reproduction. In the genusCycasthe female flower is peculiar among cycads in consisting of a terminal crown of separate leaf-like carpels several inches in length; the apical portion of each carpellary leaf may be broadly triangular in form, and deeply dissected on the margins into narrow woolly appendages like rudimentary pinnae. From the lower part of a carpel are produced several laterally placed ovules, which become bright red or orange on ripening; the bright fleshy seeds, which in some species are as large as a goose’s egg, and the tawny spreading carpels produce a pleasing combination of colour in the midst of the long dark-green fronds, which curve gracefully upwards and outwards from the summit of the columnar stem. InCycasthe stem apex, after producing a cluster of carpellary leaves, continues to elongate and produces more bud-scales, which are afterwards pushed aside as a fresh crown of fronds is developed. The young leaves ofCycasconsist of a straight rachis bearing numerous linear pinnae, traversed by a single midrib; the pinnae are circinately coiled like the leaf of a fern (fig. 3). The male flower ofCycasconforms to the type of structure characteristic of the cycads, and consists of a long cone of numerous sporophylls bearing many oval pollen-sacs on their lower faces. The type described serves as a convenient representative of its class. There are eight other living genera, which may be classified as follows:—
Classification.—A.Cycadeae.—Characterized by (a) the alternation of scale- and foliage-leaves (fig. 1) on the branched or unbranched stem; (b) the growth of the main stem through the female flower; (c) the presence of a prominent single vein in the linear pinnae; (d) the structure of the female flower, which is peculiar in not having the form of a cone, but consists of numerous independent carpels, each of which bears two or more lateral ovules. Represented by a single genus,Cycas. (Tropical Asia, Australia, &c.).
B.Zamieae.—The stem does not grow through the female flower; both male and female flowers are in the form of cones. (a)Stangerieae.—Characterized by the fern-like venation of the pinnae, which have a prominent midrib, giving off at a wide angle simple or forked and occasionally anastomosing lateral veins. A single genus,Stangeria, confined to South Africa, (b)Euzamieae.—The pinnae are traversed by several parallel veins.Bowenia, an Australian cycad, is peculiar in having bi-pinnate fronds (fig. 5). The various genera are distinguished from one another by the shape and manner of attachment of the pinnae, the form of the carpellary scales, and to some extent by anatomical characters.Encephalartos(South and Tropical Africa).—Large cones; the carpellary scales terminate in a peltate distal expansion.Macrozamia(Australia).—Similar toEncephalartosexcept in the presence of a spinous projection from the swollen distal end of the carpels.Zamia(South America, Florida, &c.).—Stem short and often divided into several columnar branches. Each carpel terminates in a peltate head.Ceratozamia(Mexico).—Similar in habit toMacrozamia, but distinguished by the presence of two horn-like spinous processes on the apex of the carpels.Microcycas(Cuba).—LikeZamia, except that the ends of the stamens are flat, while the apices of the carpels are peltate.Dioon(Mexico) (fig. 4).—Characterized by the woolly scale-leaves and carpels; the latter terminate in a thick laminar expansion of triangular form, bearing two placental cushions, on which the ovules are situated.Bowenia(Australia).—Bi-pinnate fronds; stem short and tuberous (fig. 5).
The stems of cycads are often described as unbranched; it is true that in comparison with conifers, in which the numerous branches, springing from the main stem, give a characteristic form to the tree, the tuberous or columnar stem of the CycadaceaeStem and leaf.constitutes a striking distinguishing feature. Branching, however, occurs not infrequently: inCycasthe tall stem often produces several candelabra-like arms; inZamiathe main axis may break up near the base into several cylindrical branches; in species ofDioon(fig. 4) lateral branches are occasionally produced. The South AfricanEncephalartosfrequently produces several branches. Probably the oldest example of this genus in cultivation is in the Botanic Garden of Amsterdam, its age is considered by Professor de Vries to be about two thousand years: although an accurate determination of age is impossible, there is no doubt that many cycads grow very slowly and are remarkable for longevity. The thick armour of petiole-bases enveloping the stem is a characteristic Cycadean feature; inCycasthe alternation of scale-leaves and fronds is more clearly shown than in other cycads; inEncephalartos,Dioon, &c., the persistent scale-leaf bases are almost equal in size to those of the foliage-leaves, and there is no regular alternation of zones such as characterizes some species ofCycas. Another type of stem is illustrated byStangeriaandZamia, also by a few forms ofCycas(fig. 2), in which the fronds fall off completely, leaving a comparatively smooth stem. TheCyastype of frond, except as regards the presence of a midrib in each pinna, characterizes the cycads generally, exceptBoweniaandStangeria. In the monotypic genusBoweniathe large fronds, borne singly on the short and thick stem, are bi-pinnate (fig. 5); the segments, which are broadly ovate or rhomboidal, have several forked spreading veins, and resemble the large pinnules of some species ofAdiantum. InStangeria, also a genus represented by one species (S. paradoxaof South Africa), the long and comparatively broad pinnae, with an entire or irregularly incised margin, are very fern-like, a circumstance which led Kunze to describe the plant in 1835 as a species of the fernLomaria. In rare cases the pinnae of cycads are lobed or branched: inDioon spinulosum(Central America) the margin of the segments bears numerous spinous processes; in some species ofEncephalartos,e.g.E. horridus, the lamina is deeply lobed; and in a species of the Australian genusMacrozamia,M. heteromera, the narrow pinnae are dichotomously branched almost to the base (fig. 6), and resemble the frond of some species of the fernSchizaea, or the fossil genusBaiera(Ginkgoales). An interesting species ofCycas,C. Micholitzii, has recently been described by Sir William Thiselton-Dyer from Annam, where it was collected by one of Messrs Sanders & Son’s collectors, in which the pinnae instead of being of the usual simple type aredichotomously branched as inMacrozamia heteromera. InCeratozamiathe broad petiole-base is characterized by the presence of two lateral spinous processes, suggesting stipular appendages, comparable, on a reduced scale, with the large stipules of the Marattiaceae among Ferns. The vernation varies in different genera; inCycasthe rachis is straight and the pinnae circinately coiled (fig. 3); inEncephalartos,Dioon, &c., both rachis and segments are straight; inZamiathe rachis is bent or slightly coiled, bearing straight pinnae. The young leaves arise on the stem-apex as conical protuberances with winged borders on which the pinnae appear as rounded humps, usually in basipetal order; the scale-leaves in their young condition resemble fronds, but the lamina remains undeveloped. A feature of interest in connexion with the phylogeny of cycads is the presence of long hairs clothing the scale-leaves, and forming a cap on the summit of the stem-apex or attached to the bases of petioles; on some fossil cycadean plants these outgrowths have the form of scales, and are identical in structure with the ramenta (paleae) of the majority of ferns.
The male flowers of cycads are constructed on a uniform plan, and in all cases consist of an axis bearing crowded, spirally disposed sporophylls. These are often wedge-shaped and angular; in some cases they consist of a short, thickFlower.stalk, terminating in a peltate expansion, or prolonged upwards in the form of a triangular lamina. The sporangia (pollen-sacs), which occur on the under-side of the stamens, are often arranged in more or less definite groups or sori, interspersed with hairs (paraphyses); dehiscence takes place along a line marked out by the occurrence of smaller and thinner-walled cells bounded by larger and thicker-walled elements, which form a fairly prominent cap-like “annulus” near the apex of the sporangium, not unlike the annulus characteristic of the Schizaeaceae among ferns. The sporangial wall, consisting of several layers of cells, encloses a cavity containing numerous oval spores (pollen-grains). In structure a cycadean sporangium recalls those of certain ferns (Marattiaceae, Osmundaceae and Schizaeaceae), but in the development of the spores there are certain peculiarities not met with among the Vascular Cryptogams. With the exception ofCycas, the female flowers are also in the form of cones, bearing numerous carpellary scales. InCycas revolutaandC. circinaliseach leaf-like carpel may produce several laterally attached ovules, but inC. Normanbyanathe carpel is shorter and the ovules are reduced to two; this latter type brings us nearer to the carpels ofDioon, in which the flower has the form of a cone, and the distal end of the carpels is longer and more leaf-like than in the other genera of theZamieae, which are characterized by shorter carpels with thick peltate heads bearing two ovules on the morphologically lower surface. The cones of cycads attain in some cases (e.g.Encephalartos) a considerable size, reaching a length of more than a foot. Cases have been recorded (by Thiselton-Dyer inEncephalartosand by Wieland inZamia) in which the short carpellary cone-scales exhibit a foliaceous form. It is interesting that no monstrous cycadean cone has been described in which ovuliferous and staminate appendages are borne on the same axis: in the Bennettitales (seePalaeobotany:Mesozoic) flowers were produced bearing on the same axis both androecium and gynoecium.
P, Prothallus.
A, Archegonia.
N, Nucellus.
C, Pollen-chamber.
Pt, Pollen-tube.
Pg, Pollen-grain.
G, Generative cell (second cell of pollen-tube).
The pollen-grains when mature consist of three cells, two small and one large cell; the latter grows into the pollen-tube, as in the Coniferales, and from one of the small cells two large ciliated spermatozoids are eventually produced. AMicrospores and megaspores.remarkable exception to this rule has recently been recorded by Caldwell, who found that inMicrocycas Calocomathe body-cells may be eight or even ten in number and the sperm-cells twice as numerous. One of the most important discoveries made during the latter part of the 19th century was that by Ikeno, a Japanese botanist, who first demonstrated the existence of motile male cells in the genusCycas. Similar spermatozoids were observed in some species ofZamiaby H. J. Webber, and more recent work enables us to assume that all cycads produce ciliated male gametes. Before following the growth of the pollen-grain after pollination, we will briefly describe the structure of a cycadean ovule. An ovule consists of a conical nucellus surrounded by a single integument. At an early stage of development a large cell makes its appearance in the central region of the nucellus; this increases in size and eventually forms three cells; the lowest of these grows vigorously and constitutes the megaspore (embryo-sac), which ultimately absorbs the greater part of the nucellus. The megaspore-nucleus divides repeatedly, and cells are produced from the peripheral region inwards, which eventually fill the spore-cavity with a homogeneous tissue (prothallus); some of the superficial cells at the micropylar end of the megaspore increase in size and divide by a tangential wall into two, an upper cell which gives rise to the short two-celled neck of the archegonium, and a lower cell which develops into a large egg-cell. Each megaspore may contain 2 to 6 archegonia. During the growth of the ovum nourishment is supplied from the contents of the cells immediately surrounding the egg-cell, as in the development of the ovum ofPinusand other conifers. Meanwhile the tissue in the apical region of the nucellus has been undergoing disorganization, which results in the formation of a pollen-chamber (fig. 7,C) immediately above the megaspore. Pollination in cycads has always been described as anemophilous, but according to recent observations by Pearson on South African species it seems probable that, at least in some cases, the pollen is conveyed to the ovules by animal agency. The pollen-grains find their way between the carpophylls, which at the time of pollination are slightly apart owing to the elongation of the internodes of the flower-axis, and pass into the pollen-chamber; the large cell of the pollen-grain grows out into a tube (Pt), which penetrates the nucellar tissue and often branches repeatedly; the pollen-grain itself, with the prothallus-cells, projects freely into the pollen-chamber (fig. 7). The nucleus of the outermost (second) small cell (fig. 7,G) divides, and one of the daughter-nuclei passes out of the cell, and may enter the lowest (first) small cell. The outermost cell, by the division of the remaining nucleus, produces two large spermatozoids (fig. 8,a,a). InMicrocycas16 sperm-cells are produced. In the course of division two bodies appear in the cytoplasm, and behave as centrosomes during the karyokinesis; they gradually become threadlike and coil round each daughter nucleus. This thread gives rise to a spiral ciliated band lying in a depression on the body of each spermatozoid; the large spermatozoids eventually escape from the pollen-tube, and are able to perform ciliary movements in the watery liquid which occurs between the thin papery remnant of nucellar tissue and the archegonial necks. Before fertilization a neck-canal cell is formed by the division of the ovum-nucleus. After the body of a spermatozoid has coalesced with the egg-nucleus the latter divides repeatedly and forms a mass of tissue which grows more vigorously in the lower part of the fertilized ovum, and extends upwards towards the apex of the ovum as a peripheral layer of parenchyma surrounding a central space. By further growth this tissue gives rise to a proembryo, which consists, at the micropylar end, of a sac; the tissue at the chalazal end grows into a long and tangled suspensor, terminating in a mass of cells, which is eventually differentiated into a radicle, plumule and two cotyledons. In the ripe seed the integument assumes the form of a fleshy envelope, succeeded internally by a hard woody shell, internal to which is a thin papery membrane—the apical portion of the nucellus—which is easily dissected out as a conical cap covering the apex of the endosperm. A thorough examination of cycadean seeds has recently been made by Miss Stopes, more particularly with a view to a comparison of their vascular supply with that in Palaeozoic gymnospermous seeds (Flora, 1904). The first leaves borne on the seedling axis are often scale-like, and these are followed by two or more larger laminae, which foreshadow the pinnae of the adult frond.
pd, Periderm in leaf-bases.
lt, Leaf-traces in cortex.
ph, Phloem.
x, Xylem.
m, Medullary bundles.
c, Cortical bundles.
The anatomical structure of the vegetative organs of recent cycads is of special interest as affording important evidence of relationship with extinct types, and with other groups of recent plants. Brongniart, whoAnatomy.was the first to investigate in detail the anatomy of a cycadean stem, recognized an agreement, as regards the secondary wood, with Dicotyledons and Gymnosperms, rather than with Monocotyledons. He drew attention also to certain structural similarities betweenCycasandGinkgo. The main anatomical features of a cycad stem may be summarized as follows: the centre is occupied by a large parenchymatous pith traversed by numerous secretory canals, and in some genera by cauline vascular bundles (e.g.EncephalartosandMacrozamia). In addition to these cauline strands (confined to the stem and not connected with the leaves), collateral bundles are often met with in the pith, which form the vascular supply of terminal flowers borne at intervals on the apex of the stem. These latter bundles may be seen in sections of old stems to pursue a more or less horizontal course, passing outwards through the main woody cylinder. This lateral course is due to the more vigorous growth of the axillary branch formed near the base of each flower, which is a terminal structure, and, except in the female flower ofCycas, puts a limit to the apical growth of the stem. The vigorous lateral branch therefore continues the line of the main axis. The pith is encircled by a cylinder of secondary wood, consisting of single or multiple radial rows of tracheids separated by broad medullary rays composed of large parenchymatous cells; the tracheids bear numerous borderedpits on the radial walls. The large medullary rays give to the wood a characteristic parenchymatous or lax appearance, which is in marked contrast to the more compact wood of a conifer. The protoxylem-elements are situated at the extreme inner edge of the secondary wood, and may occur as small groups of narrow, spirally-pitted elements scattered among the parenchyma which abuts on the main mass of wood. Short and reticulately-pitted tracheal cells, similar to tracheids, often occur in the circummedullary region of cycadean stems. In an old stem ofCycas,EncephalartosorMacrozamiathe secondary wood consists of several rather unevenly concentric zones, while in some other genera it forms a continuous mass as in conifers and normal dicotyledons. These concentric rings of secondary xylem and phloem (fig. 9) afford a characteristic cycadean feature. After the cambium has been active for some time producing secondary xylem and phloem, the latter consisting of sieve-tubes, phloem-parenchyma and frequently thick-walled fibres, a second cambium is developed in the pericycle; this produces a second vascular zone, which is in turn followed by a third cambium, and so on, until several hollow cylinders are developed. It has been recently shown that several cambium-zones may remain in a state of activity, so that the formation of a new cambium does not necessarily mark a cessation of growth in the more internal meristematic rings. It occasionally happens that groups of xylem and phloem are developed internally to some of the vascular rings; these are characterized by an inverse orientation of the tissues, the xylem being centrifugal and the phloem centripetal in its development. The broad cortical region, which contains many secretory canals, is traversed by numerous vascular bundles (fig. 9,c) some of which pursue a more or less vertical course, and by frequent anastomoses with one another form a loose reticulum of vascular strands; others are leaf-traces on their way from the stele of the stem to the leaves. Most of these cortical bundles are collateral in structure, but in some the xylem and phloem are concentrically arranged; the secondary origin of these bundles from procambium-strands was described by Mettenius in his classical paper of 1860. During the increase in thickness of a cycadean stem successive layers of cork-tissue are formed by phellogens in the persistent bases of leaves (fig. 9,pd), which increase in size to adapt themselves to the growth of the vascular zones. The leaf-traces of cycads are remarkable both on account of their course and their anatomy. In a transverse section of a stem (fig. 9) one sees some vascular bundles following a horizontal or slightly oblique course in the cortex, stretching for a longer or shorter distance in a direction concentric with the woody cylinder. From each leaf-base two main bundles spread right and left through the cortex of the stem (fig. 9,lt), and as they curve gradually towards the vascular ring they present the appearance of two rather flat ogee curves, usually spoken of as the leaf-trace girdles (fig. 9,lt). The distal ends of these girdles give off several branches, which traverse the petiole and rachis as numerous collateral bundles. The complicated girdle-like course is characteristic of the leaf-traces of most recent cycads, but in some cases,e.g.inZamia floridana, the traces are described by Wieland in his recent monograph on American fossil cycads (Carnegie Institution Publications, 1906) as possessing a more direct course similar to that in Mesozoic genera. A leaf-trace, as it passes through the cortex, has a collateral structure, the protoxylem being situated at the inner edge of the xylem; when it reaches the leaf-base the position of the spiral tracheids is gradually altered, and the endarch arrangement (protoxylem internal) gives place to a mesarch structure (protoxylem more or less central and not on the edge of the xylem strand). In a bundle examined in the basal portion of a leaf the bulk of the xylem is found to be centrifugal in position, but internally to the protoxylem there is a group of centripetal tracheids; higher up in the petiole the xylem is mainly centripetal, the centrifugal wood being represented by a small arc of tracheids external to the protoxylem and separated from it by a few parenchymatous elements. Finally, in the pinnae of the frond the centrifugal xylem may disappear, the protoxylem being now exarch in position and abutting on the phloem. Similarly in the sporophylls of some cycads the bundles are endarch near the base and mesarch near the distal end of the stamen or carpel. The vascular system of cycadean seedlings presents some features worthy of note; centripetal xylem occurs in the cotyledonary bundles associated with transfusion-tracheids. The bundles from the cotyledons pursue a direct course to the stele of the main axis, and do not assume the girdle-form characteristic of the adult plant. This is of interest from the point of view of the comparison of recent cycads with extinct species (Bennettites), in which the leaf-traces follow a much more direct course than in modern cycads. The mesarch structure of the leaf-bundles is met with in a less pronounced form in the flower peduncles of some cycads. This fact is of importance as showing that the type of vascular structure, which characterized the stems of many Palaeozoic genera, has not entirely disappeared from the stems of modern cycads; but the mesarch bundle is now confined to the leaves and peduncles. The roots of some cycadsRoots.resemble the stems in producing several cambium-rings; they possess 2 to 8 protoxylem-groups, and are characterized by a broad pericyclic zone. A common phenomenon in cycads is the production of roots which grow upwards (apogeotropic), and appear as coralline branched structures above the level of the ground; some of the cortical cells of these roots are hypertrophied, and contain numerous filaments of blue-green Algae (Nostocaceae), which live as endoparasites in the cell-cavities.
Ginkgoales.—This class-designation has been recently proposed to give emphasis to the isolated position of the genusGinkgo(Salisburia) among the Gymnosperms.Ginkgo biloba, the maidenhair tree, has usually been placed by botanists in the Taxeae in the neighbourhood of the yew (Taxus), but the proposal by Eichler in 1852 to institute a special family, theSalisburieae, indicated a recognition of the existence of special characteristics which distinguish the genus from other members of the Coniferae. The discovery by the Japanese botanist Hirase of the development of ciliated spermatozoids in the pollen-tube ofGinkgo, in place of the non-motile male cells of typical conifers, served as a cogent argument in favour of separating the genus from the Coniferales and placing it in a class of its own. In 1712 Kaempfer published a drawing of a Japanese tree, which he described under the nameGinkgo; this term was adopted in 1771 by Linnaeus, who spoke of Kaempfer’s plant asGinkgo biloba. In 1797 Smith proposed to use the nameSalisburia adiantifoliain preference to the “uncouth” genusGinkgoand “incorrect” specific termbiloba. Both names are still in common use. On account of the resemblance of the leaves to those of some species ofAdiantum, the appellation maidenhair tree has long been given toGinkgo biloba.Ginkgois of special interest on account of its isolated position among existing plants, its restricted geographical distribution, and its great antiquity (seePalaeobotany:Mesozoic). This solitary survivor of an ancient stock is almost extinct, but a few old and presumably wild trees are recorded by travellers in parts of China.Ginkgois common as a sacred tree in the gardens of temples in the Far East, and often cultivated in North America and Europe.Ginkgo biloba, which may reach a height of over 30 metres, forms a tree of pyramidal shape with a smooth grey bark. The leaves (figs. 10 and 11) have a long, slender petiole terminating in a fan-shaped lamina, which may be entire, divided by a median incision into two wedge-shaped lobes, or subdivided into several narrow segments. The venation is like that of many ferns,e.g.Adiantum; the lowest vein in each half of the lamina follows a course parallel to the edge, and gives off numerous branches, which fork repeatedly as they spread in a palmate manner towards the leaf margin. The foliage-leaves occur either scattered on long shoots of unlimited growth, or at the apex of short shoots (spurs), which may eventually elongate into long shoots.
p, Pollen-tube (proximal end).
c, Pollen-chamber.
e, Upward prolongation of megaspore.
a, Archegonia.
Pg, Pollen-grain.
Ex, Exine.
The flowers are dioecious. The male flowers (fig. 12), borne in the axil of scale-leaves, consist of a stalked central axis bearing loosely disposed stamens; each stamen consists of a slender filament terminating in a small apical scale, which bearsFlowers.usually two, but not infrequently three or four pollen-sacs (fig. 12,C). The axis of the flower is a shoot bearing leaves in the form of stamens. A mature pollen-grain contains a prothallus of 3 to 5 cells (Fig. 13, Pg); the exine extends over two-thirds of the circumference, leaving a thin portion of the wall, which on collapsing produces a longitudinal groove similar to the median depression on the pollen-grain of a cycad. The ordinary type of female flower has the form of a long, naked peduncle bearing a single ovule on either side of the apex (fig. 12), the base of each being enclosed by a small, collar-like rim, the nature of which has been variously interpreted. A young ovule consists of a conical nucellus surrounded by a single integument terminating as a two-lipped micropyle. A large pollen-chamber occupies the apex of the nucellus; immediately below this, two or more archegonia (fig. 13,a) are developed in the upper region of the megaspore, each consisting of a large egg-cell surmounted by two neck-cells and a canal-cell which is cut off shortly before fertilization. After the entrance of the pollen-grain the pollen-chamber becomes roofed over by a blunt protuberance of nucellar tissue. The megaspore (embryo-sac) continues to grow after pollination until the greater part of the nucellus is gradually destroyed; it also gives rise to a vertical outgrowth, which projects from the apex of the megaspore as a short, thick column (fig. 13,e) supporting the remains of the nucellar tissue which forms the roof of the pollen-chamber (fig. 13,c). Surrounding the pitted wall of the ovum there is a definite layer of large cells, no doubt representing a tapetum, which, as in cycads and conifers, plays an important part in nourishing the growing egg-cell. The endosperm detached from a largeGinkgoovule after fertilization bears a close resemblance to that of a cycad; the apex is occupied by a depression, on the floor of which two small holes mark the position of the archegonia, and the outgrowth from the megaspore apex projects from the centre as a short peg. After pollination the pollen-tube grows into the nucellar tissue, as in cycads, and the pollen-grain itself (fig. 13,Pg) hangs down into the pollen-chamber; two large spirally ciliated spermatozoids are produced, their manner of development agreeing very closely with that of the corresponding cells inCycasandZamia. After fertilization the ovum-nucleus divides and cell-formation proceeds rapidly, especially in the lower part of the ovum, in which the cotyledon and axis of the embryo are differentiated; the long, tangled suspensor of the cycadean embryo is not found inGinkgo. It is often stated that fertilization occurs after the ovules have fallen, but it has been demonstrated by Hirase that this occurs while the ovules are still attached to the tree. The ripe seed, which grows as large as a rather small plum, is enclosed by a thick, fleshy envelope covering a hard woody shell with two or rarely three longitudinal keels. A papery remnant of nucellus lines the inner face of the woody shell, and, as in cycadean seeds, the apical portion is readily separated as a cap covering the summit of the endosperm.
The morphology of the female flowers has been variously interpreted by botanists; the peduncle bearing the ovules has been described as homologous with the petiole of a foliage-leaf and as a shoot-structure, the collar-like envelope at the base of the ovules being referred to as a second integument or arillus, or as the representative of a carpel. The evidence afforded by normal and abnormal flowers appears to be in favour of the following interpretation: The peduncle is a shoot bearing two or more carpels. Each ovule is enclosed at the base by an envelope or collar homologous with the lamina of a leaf; the fleshy and hard coats of the nucellus constitute a single integument. The stalk of an ovule, considerably reduced in normal flowers and much larger in some abnormal flowers, is homologous with a leaf-stalk, with which it agrees in the structure and number of vascular bundles. The facts on which this description is based are derived partly from anatomical evidence, and in part from an account given by a Japanese botanist, Fujii, of several abnormal female flowers; in some cases the collar at the base of an ovule, often described as an arillus, is found to pass gradually into the lamina of a leaf bearing marginal ovules (fig. 14,B). The occurrence of more than two ovules on one peduncle is by no means rare; a particularly striking example is described by Fujii, in which an unusually thick peduncle bearing several stalked ovules terminates in a scaly bud (fig. 14,A,b). The frequent occurrence of more than two pollen-sacs and the equally common occurrence of additional ovules have been regarded by some authors as evidence in favour of the view that ancestral types normally possessed a greater number of these organs than are usually found in the recent species. ThisAnatomy.view receives support from fossil evidence. Close to the apex of a shoot the vascular bundles of a leaf make their appearance as double strands, and the leaf-traces in the upper part of a shoot have the form of distinct bundles, which in the older part of the shoot form a continuous ring. Each double leaf-trace passes through four internodes before becoming a part of the stele; the double nature of the trace is a characteristic feature. Secretory sacs occur abundantly in the leaf-lamina, where they appear as short lines between the veins; they are abundant also in the cortex and pith of the shoot, in the fleshy integument of the ovule, and elsewhere. The secondary wood of the shoot and root conforms in the main to the coniferous type; in the short shoots the greater breadth of the medullary rays in the more internal part of the xylem recalls the cycadean type. The secondary phloem contains numerous thick-walled fibres, parenchymatous cells, and large sieve-tubes with plates on the radial walls; swollen parenchymatous cells containing crystals are commonly met with in the cortex, pith and medullary-ray tissues. The wood consists of tracheids, with circular bordered pits on their radial walls, and in the late summer wood pits are unusually abundant on the tangential walls. A point of anatomical interest is the occurrence in the vascular bundles of the cotyledons, scale-leaves, and elsewhere of a few centripetally developed tracheids, which give to the xylem-strands a mesarch structure such as characterizes the foliar bundles of cycads. The root is diarch in structure, but additional protoxylem-strands may be present at the base of the main root; the pericycle consists of several layers of cells.
This is not the place to discuss in detail the past history ofGinkgo(seePalaeobotany:Mesozoic). Among Palaeozoic genera there are some which bear a close resemblance to the recent type in the form of the leaves; and petrified Palaeozoic seeds,Geological history.almost identical with those of the maidenhair tree, have been described from French and English localities. During the Triassic and Jurassic periods the genusBaiera—no doubt a representative of the Ginkgoales—was widely spread throughout Europe and in other regions;Ginkgoitself occurs abundantly in Mesozoic and Tertiary rocks, and was a common plant in the Arctic regions as elsewhere during the Jurassic and Lower Cretaceous periods. Some unusually perfectGinkgoleaves have been found in the Eocene leaf-beds between the lava-flows exposed in the cliffs of Mull (fig. 11). From an evolutionary point of view, it is of interest to note the occurrence of filicinean and cycadean characters in the maidenhair tree. The leaves at once invite a comparison with ferns; the numerous long hairs which form a delicate woolly covering on young leaves recall the hairs of certain ferns, but agree more closely with the long filamentous hairs of recent cycads. The spermatozoids constitute the most striking link with both cycads and ferns. The structure of the seed, the presence of two neck-cells in the archegonia, the late development of the embryo, the partially-fused cotyledons and certain anatomical characters, are features common toGinkgoand the cycads. The maidenhair tree is one of the most interesting survivals from the past; it represents a type which, in the Palaeozoic era, may have been merged into the extinct class Cordaitales. Through the succeeding ages the Ginkgoales were represented by numerous forms, which gradually became more restricted in their distribution and fewer in number during the Cretaceous and Tertiary periods, terminating at the present day in one solitary survivor.
Coniferales.—Trees and shrubs characterized by a copious branching of the stem and frequently by a regular pyramidal form. Leaves simple, small, linear or short and scale-like, usually persisting for more than one year. Flowers monoecious or dioecious, unisexual, without a perianth, often in the form of cones, but never terminal on the main stem.
The plants usually included in the Coniferae constitute a less homogeneous class than the Cycadaceae. Some authors use the term Coniferae in a restricted sense as including those genera which have the female flowers in the form of cones,External features.the other genera, characterized by flowers of a different type, being placed in the Taxaceae, and often spoken of as Taxads.In order to avoid confusion in the use of the term Coniferae, we may adopt as a class-designation the name Coniferales, including both the Coniferae—using the term in a restricted sense—and the Taxaceae. The most striking characteristic of the majority of the Coniferales is the regular manner of the monopodial branching and the pyramidal shape.Araucaria imbricata, the Monkey-puzzle tree,A. excelsa, the Norfolk Island pine, many pines and firs, cedars and other genera illustrate the pyramidal form. The mammoth redwood tree of California,Sequoia(Wellingtonia)gigantea, which represents the tallest Gymnosperm, is a good example of the regular tapering main stem and narrow pyramidal form. The cypresses afford instances of tall and narrow trees similar in habit to Lombardy poplars. The common cypress (Cupressus sempervirens), as found wild in the mountains of Crete and Cyprus, is characterized by long and spreading branches, which give it a cedar-like habit. A pendulous or weeping habit is assumed by some conifers,e.g.Picea excelsavar.virgatarepresents a form in which the main branches attain a considerable horizontal extension, and trail themselves like snakes along the ground. Certain species ofPinus, the yews (Taxus) and some other genera grow as bushes, which in place of a main mast-like stem possess several repeatedly-branched leading shoots. The unfavourable conditions in Arctic regions have produced a dwarf form, in which the main shoots grow close to the ground. Artificially induced dwarfed plants ofPinus,Cupressus,Sciadopitys(umbrella pine) and other genera are commonly cultivated by the Japanese. The dying off of older branches and the vigorous growth of shoots nearer the apex of the stem produce a form of tree illustrated by the stone pine of the Mediterranean region (Pinus Pinea), which Turner has rendered familiar in his “Childe Harold’s Pilgrimage” and other pictures of Italian scenery. Conifers are not infrequently seen in which a lateral branch has bent sharply upwards to take the place of the injured main trunk. An upward tendency of all the main lateral branches, known as fastigiation, is common in some species, producing well-marked varieties,e.g.Cephalotaxus pedunculatavar.fastigiata; this fastigiate habit may arise as a sport on a tree with spreading branches. Another departure from the normal is that in which the juvenile or seedling form of shoot persists in the adult tree; the numerous coniferous plants known as species ofRetinosporaare examples of this. The nameRetinospora, therefore, does not stand for a true genus, but denotes persistent young forms ofJuniperus,Thuja,Cupressus, &c., in which the small scaly leaves of ordinary species are replaced by the slender, needle-like leaves, which stand out more or less at right angles from the branches. The flat branchlets ofCupressus,Thuja(arbor vitae),Thujopsis dolabrata(Japanese arbor vitae) are characteristic of certain types of conifers; in some cases the horizontal extension of the branches induces a dorsiventral structure. A characteristic feature of the genusAgathis(Dammara) the Kauri pine of New Zealand, is the deciduous habit of the branches; these become detached from the main trunk leaving a well-defined absciss-surface, which appears as a depressed circular scar on the stem. A new genus of conifers,Taiwania, has recently been described from the island of Formosa; it is said to agree in habit with the JapaneseCryptomeria, but the cones appear to have a structure which distinguishes them from those of any other genus.
With a few exceptions conifers are evergreen, and retain the leaves for several years (10 years inAraucaria imbricata, 8 to 10 inPicea excelsa, 5 inTaxus baccata; inPinusthe needles usually fall in October of their third year). The larch (Larix)Leaves.sheds its leaves in the autumn, in the Chinese larch (Pseudolarix Kaempferi) the leaves turn a bright yellow colour before falling. In the swamp cypress (Taxodium distichum) the tree assumes a rich brown colour in the autumn, and sheds its leaves together with the branchlets which bear them; deciduous branches occur also in some other species,e.g.Sequoia sempervirens(redwood),Thuja occidentalis, &c. The leaves of conifers are characterized by their small size,e.g.the needle-form represented byPinus,Cedrus,Larix, &c., the linear flat or angular leaves, appressed to the branches, ofThuja,Cupressus,Libocedrus, &c. The flat and comparatively broad leaves ofAraucaria imbricata,A. Bidwillii, and some species of the southern genusPodocarpusare traversed by several parallel veins, as are also the still larger leaves ofAgathis, which may reach a length of several inches. In addition to the foliage-leaves several genera also possess scale-leaves of various kinds, represented by bud-scales inPinus,Picea, &c., which frequently persist for a time at the base of a young shoot which has pushed its way through the yielding cap of protecting scales, while in some conifers the bud-scales adhere together, and after being torn near the base are carried up by the growing axis as a thin brown cap. The cypresses, araucarias and some other genera have no true bud-scales; in some species,e.g.Araucaria Bidwillii, the occurrence of small foliage-leaves, which have functioned as bud-scales, at intervals on the shoots affords a measure of seasonal growth. The occurrence of long and short shoots is a characteristic feature of many conifers. InPinusthe needles occur in pairs, or in clusters of 3 or 5 at the apex of a small and inconspicuous short shoot of limited growth (spur), which is enclosed at its base by a few scale-leaves, and borne on a branch of unlimited growth in the axil of a scale-leaf. In the CalifornianPinus monophyllaeach spur bears usually one needle, but two are not uncommon; it would seem that rudiments of two needles are always produced, but, as a rule, only one develops into a needle. InSciadopityssimilar spurs occur, each bearing a single needle, which in its grooved surface and in the possession of a double vascular bundle bears traces of an origin from two needle-leaves. A peculiarity of these leaves is the inverse orientation of the vascular tissue; each of the two veins has its phloem next the upper and the xylem towards the lower surface of the leaf; this unusual position of the xylem and phloem may be explained by regarding the needle ofSciadopitysas being composed of a pair of leaves borne on a short axillary shoot and fused by their margins (fig. 15, A). Long and short shoots occur also inCedrusandLarix, but in these genera the spurs are longer and stouter, and are not shed with the leaves; this kind of short shoot, by accelerated apical growth, often passes into the condition of a long shoot on which the leaves are scattered and separated by comparatively long internodes, instead of being crowded into tufts such as are borne on the ends of the spurs. In the genusPhyllocladus(New Zealand, &c.) there are no green foliage-leaves, but in their place flattened branches (phylloclades) borne in the axils of small scale-leaves. The cotyledons are often two in number, but sometimes (e.g.Pinus) as many as fifteen; these leaves are usually succeeded by foliage-leaves in the form of delicate spreading needles, and these primordial leaves are followed, sooner or later, by the adult type of leaf, except in Retinosporas, which retain the juvenile foliage. In addition to the first foliage-leaves and the adult type of leaf, there are often produced leaves which are intermediate both in shape and structure between the seedling and adult foliage. Dimorphism or heterophylly is fairly common. One of the best known examples is the Chinese juniper (Juniperus chinensis), in which branches with spinous leaves, longer and more spreading than the ordinary adult leaf, are often found associated with the normal type of branch. In some cases,e.g.Sequoia sempervirens, the fertile branches bear leaves which are less spreading than those on the vegetative shoots. Certain species of the southern hemisphere genusDacrydiumafford particularly striking instances of heterophylly,e.g.D. Kirkiiof New Zealand, in which some branches bear small and appressed leaves, while in others the leaves are much longer and more spreading. A well-known fossil conifer from Triassic strata—Voltzia heterophylla—also illustrates a marked dissimilarity in the leaves of the same shoot. The variation in leaf-form and the tendency of leaves to arrange themselves in various ways on different branches of the same plant are features which it is important to bear in mind in the identification of fossil conifers. In this connexion we may note the striking resemblance between some of the New Zealand Alpine Veronicas,e.g.Veronica Hectori,V. cupressoides, &c. (alsoPolycladus cupressinus, a Composite), and some of the cypresses and other conifers with small appressed leaves. The long linear leaves of some species ofPodocarpus, in which the lamina is traversed by a single vein, recall the pinnae of Cycas; the branches of some Dacrydiums and other forms closely resemble those of lycopods; these superficial resemblances, both between different genera of conifers and between conifers and other plants, coupled with the usual occurrence of fossil coniferous twigs without cones attached to them, render the determination of extinct types a very unsatisfactory and frequently an impossible task.
A typical male flower consists of a central axis bearing numerous spirally-arranged sporophylls (stamens), each of which consists of a slender stalk (filament) terminating distally in a more or less prominent knob or triangular scale, and bearingFlowers.two or more pollen-sacs (microsporangia) on its lower surface. The pollen-grains of some genera (e.g.Pinus) are furnished with bladder-like extensions of the outer wall, which serve as aids to wind-dispersal. The stamens ofAraucariaandAgathisare peculiar in bearing several long, and narrow free pollen-sacs; these may be compared with the sporangiophores of the horsetails (Equisetum); inTaxus(yew) the filament is attached to the centre of a large circular distal expansion, which bears several pollen-sacs on its under surface. In the conifers proper the female reproductive organs have the form of cones, which may be styled flowers or inflorescences according to different interpretations of their morphology. In the Taxaceae the flowers have a simpler structure. The female flowers of theAbietineaemay be taken as representing a common type. A pine cone reaches maturity in two years; a single year suffices for the full development inLarixand several other genera. The axis of the cone bears numerous spirally disposed flat scales (cone-scales), each of which, if examined in a young cone, is found to be double, and to consist of a lower and an upper portion. The latter is a thin flat scale bearing a median ridge or keel (e.g.Abies), on each side of which is situated an inverted ovule, consisting of a nucellus surrounded by a single integument. As the cone grows in size and becomes woody the lower half of the cone-scale, which we may call the carpellary scale, may remain small, and is so far outgrown by the upper half (seminiferous scale) that it is hardly recognizable in the mature cone. In many species ofAbies(e.g.Abies pectinata, &c.) the ripe cone differs from those ofPinus,PiceaandCedrusin the large size of the carpellary scales, which project as conspicuous thin appendages beyond the distal margins of the broader and more woody seminiferous scales; the long carpellary scale is a prominent feature also in the cone of the Douglas pine (Pseudotsuga Douglasii). The female flowers (cones) vary considerably in size; the largest are the more or less spherical cones ofAraucaria—a single cone ofA. imbricatamay produce as many as 300 seeds, one seed to each fertile cone-scale—and the long pendentcones, 1 to 2 ft. in length, of the sugar pine of California (Pinus Lambertiana) and other species. Smaller cones, less than an inch long, occur in the larch,Athrotaxis(Tasmania),Fitzroya(Patagonia and Tasmania), &c. In theTaxodieaeandAraucarieaethe cones are similar in appearance to those of theAbietineae, but they differ in the fact that the scales appear to be single, even in the young condition; each cone-scale in a genus of theTaxodiinae(Sequoia, &c.) bears several seeds, while in theAraucariinae(AraucariaandAgathis) each scale has one seed. TheCupressineaehave cones composed of a few scales arranged in alternate whorls; each scale bears two or more seeds, and shows no external sign of being composed of two distinct portions. In the junipers the scales become fleshy as the seeds ripen, and the individual scales fuse together in the form of a berry. The female flowers of the Taxaceae assume another form; inMicrocachrys(Tasmania) the reproductive structures are spirally disposed, and form small globular cones made up of red fleshy scales, to each of which is attached a single ovule enclosed by an integument and partially invested by an arillus; inDacrydiumthe carpellary leaves are very similar to the foliage leaves—each bears one ovule with two integuments, the outer of which constitutes an arillus. Finally in the yew, as a type of the family Taxeae, the ovules occur singly at the apex of a lateral branch, enclosed when ripe by a conspicuous red or yellow fleshy arillus, which serves as an attraction to animals, and thus aids in the dispersal of the seeds.
Fig. 15.—Diagrammatic treatment of:
A, Double needle ofSciadopitys(a,a, leaves;b, shoot;Br, bract).
B, seminiferous scale as leaf of axillary shoot (b, shoot;Sc, seminiferous scale;Br, bract).
C, seminiferous scale as fused pair of leaves (l1,l2,l3, first, second and third leaves;b, shoot;Br, bract).
D, cone-scale ofAraucaria(n, nucellus;i, integument;x, xylem).
It is important to draw attention to some structural features exhibited by certain cone-scales, in which there is no external sign indicative of the presence of a carpellary and a seminiferous scale. InAraucaria Cookiiand some allied species eachMorphology of female flower.scale has a small pointed projection from its upper face near the distal end; the scales ofCunninghamia(China) are characterized by a somewhat ragged membranous projection extending across the upper face between the seeds and the distal end of the scale; in the scales ofAthrotaxis(Tasmania) a prominent rounded ridge occupies a corresponding position. These projections and ridges may be homologous with the seminiferous scale of the pines, firs, cedars, &c. The simplest interpretation of the cone of theAbietineaeis that which regards it as a flower consisting of an axis bearing several open carpels, which in the adult cone may be large and prominent or very small, the scale bearing the ovules being regarded as a placental outgrowth from the flat and open carpel. InAraucariathe cone-scale is regarded as consisting of a flat carpel, of which the placenta has not grown out into the scale-like structure. The seminiferous scale ofPinus, &c., is also spoken of sometimes as a ligular outgrowth from the carpellary leaf. Robert Brown was the first to give a clear description of the morphology of the Abietineous cone in which carpels bear naked ovules; he recognized gymnospermy as an important distinguishing feature in conifers as well as in cycads. Another view is to regard the cone as an inflorescence, each carpellary scale being a bract bearing in its axil a shoot the axis of which has not been developed; the seminiferous scale is believed to represent either a single leaf or a fused pair of leaves belonging to the partially suppressed axillary shoot. In 1869 van Tieghem laid stress on anatomical evidence as a key to the morphology of the cone-scales; he drew attention to the fact that the collateral vascular bundles of the seminiferous scale are inversely orientated as compared with those of the carpellary scale; in the latter the xylem of each bundle is next the upper surface, while in the seminiferous scale the phloem occupies that position. The conclusion drawn from this was that the seminiferous scale (fig. 15, B,Sc) is the first and only leaf of an axillary shoot (b) borne on that side of the shoot, the axis of which is suppressed, opposite the subtending bract (fig. 15, A, B, C,Br). Another view is to apply to the seminiferous scale an explanation similar to that suggested by von Mohl in the case of the double needle ofSciadopitys, and to consider the seed-bearing scale as being made up of a pair of leaves (fig. 15, A,a,a) of an axillary shoot (b) fused into one by their posterior margins (fig. 15, A). The latter view receives support from abnormal cones in which carpellary scales subtend axillary shoots, of which the first two leaves (fig. 15, C,l1,l1) are often harder and browner than the others; forms have been described transitional between axillary shoots, in which the leaves are separate, and others in which two of the leaves are more or less completely fused. In a young cone the seminiferous scale appears as a hump of tissue at the base or in the axil of the carpellary scale, but Celakovský, a strong supporter of the axillary-bud theory, attaches little or no importance to this kind of evidence, regarding the present manner of development as being merely an example of a short cut adopted in the course of evolution, and replacing the original production of a branch in the axil of each carpellary scale. Eichler, one of the chief supporters of the simpler view, does not recognize in the inverse orientation of the vascular bundles an argument in support of the axillary-bud theory, but points out that the seminiferous scale, being an outgrowth from the surface of the carpellary scale, would, like outgrowths from an ordinary leaf, naturally have its bundles inversely orientated. In such cone-scales as show little or no external indication of being double in origin,e.g.Araucaria(fig. 15, D) Sequoia, &c., there are always two sets of bundles; the upper set, having the phloem uppermost, as in the seminiferous scale ofAbiesorPinus, are regarded as belonging to the outgrowth from the carpellary scale and specially developed to supply the ovules. Monstrous cones are fairly common; these in some instances lend support to the axillary-bud theory, and it has been said that this theory owes its existence to evidence furnished by abnormal cones. It is difficult to estimate the value of abnormalities as evidence bearing on morphological interpretation; the chief danger lies perhaps in attaching undue weight to them, but there is also a risk of minimizing their importance. Monstrosities at least demonstrate possible lines of development, but when the abnormal forms of growth in various directions are fairly evenly balanced, trustworthy deductions become difficult. The occurrence of buds in the axils of carpellary scales may, however, simply mean that buds, which are usually undeveloped in the axils of sporophylls, occasionally afford evidence of their existence. Some monstrous cones lend no support to the axillary-bud theory. InLarixthe axis of the cone often continues its growth; similarly inCephalotaxusthe cones are often proliferous. (In rare cases the proliferated portion produces male flowers in the leaf-axils.) InLarixthe carpellary scale may become leafy, and the seminiferous scale may disappear. Androgynous cones may be produced, as in the cone ofPinus rigida(fig. 16), in which the lower part bears stamens and the upper portion carpellary and seminiferous scales. An interesting case has been figured by Masters, in which scales of a cone ofCupressus Lawsonianabear ovules on the upper surface and stamens on the lower face. One argument that has been adduced in support of the axillary bud theory is derived from the Palaeozoic typeCordaites, in which each ovule occurs on an axis borne in the axil of a bract. The whole question is still unsolved, and perhaps insoluble. It may be that the interpretation of the female cone of theAbietineaeas an inflorescence, which finds favour with many botanists, cannot be applied to the cones ofAgathisandAraucaria. Without expressing any decided opinion as to the morphology of the double cone-scale of theAbietineae, preference may be felt in favour of regarding the cone-scale of theAraucarieaeas a simple carpellary leaf bearing a single ovule. A discussion of this question may be found in a paper on theAraucarieaeby Seward and Ford, published in the Transactions of the Royal Society of London (1906).Cordaitesis an extinct type which in certain respects resemblesGinkgo, cycads and theAraucarieae, but its agreement with true conifers is probably too remote to justify our attributing much weight to the bearing of the morphology of its female flowers on the interpretation of that of the Coniferae. The greater simplicity of the Eichler theory may prejudice us in its favour; but, on the other hand, the arguments advanced in favour of the axillary-bud theories are perhaps not sufficiently cogent to lead us to accept an explanation based chiefly on the uncertain evidence of monstrosities.
A pollen-grain when first formed from its mother-cell consists of a single cell; in this condition it may be carried to the nucellus of the ovule (e.g.Taxus,Cupressus, &c.), or more usually (Pinus,Larix, &c.) it reaches maturity before the dehiscenceMicro-spores and megaspores.of the microsporangium. The nucleus of the microspore divides and gives rise to a small cell within the large cell, a second small cell is then produced; this is the structure of the ripe pollen-grain in some conifers (Taxus, &c.). The large cell grows out as a pollen-tube; the second of the two small cells (body-cell) wanders into the tube, followed by the nucleus of the first small cell (stalk-cell). InTaxusthe body-cell eventually divides into two, in which the products of division are of unequal size, the larger constituting the male generative cell, which fuses with the nucleus of the egg-cell. InJuniperusthe products of division of thebody-cell are equal, and both function as male generative cells. In theAbietineaecell-formation in the pollen-grain is carried farther. Three small cells occur inside the cavity of the microspore; two of them collapse and the third divides into two, forming a stalk-cell and a larger body-cell. The latter ultimately divides in the apex of the pollen-tube into two non-motile generative cells. Evidence has lately been adduced of the existence of numerous nuclei in the pollen-tubes of theAraucarieae, and it seems probable that in this as in several other respects this family is distinguished from other members of the Coniferales. The precise method of fertilization in the Scots Pine was followed by V. H. Blackman, who also succeeded in showing that the nuclei of the sporophyte generation contain twice as many chromosomes as the nuclei of the gametophyte. Other observers have in recent years demonstrated a similar relation in other genera between the number of chromosomes in the nuclei of the two generations. The ovule is usually surrounded by one integument, which projects beyond the tip of the nucellus as a wide-open lobed funnel, which at the time of pollination folds inwards, and so assists in bringing the pollen-grains on to the nucellus. In some conifers (e.g.Taxus,Cephalotaxus,Dacrydium, &c.) the ordinary integument is partially enclosed by an arillus or second integument. It is held by some botanists (Celakovský) that the seminiferous scale of theAbietineaeis homologous with the arillus or second integument of the Taxaceae, but this view is too strained to gain general acceptance. InAraucariaandSaxegothaeathe nucellus itself projects beyond the open micropyle and receives the pollen-grains direct. During the growth of the cell which forms the megaspore the greater part of the nucellus is absorbed, except the apical portion, which persists as a cone above the megaspore; the partial disorganization of some of the cells in the centre of the nucellar cone forms an irregular cavity, which may be compared with the larger pollen-chamber ofGinkgoand the cycads. In each ovule one megaspore comes to maturity, but, exceptionally, two may be present (e.g.Pinus sylvestris). It has been shown by Lawson that inSequoia sempervirens(Annals of Botany, 1904) and by other workers in the genera that several megaspores may attain a fairly large size in one prothallus. The megaspore becomes filled with tissue (prothallus), and from some of the superficial cells archegonia are produced, usually three to five in number, but in rare cases ten to twenty or even sixty may be present. In the genusSequoiathere may be as many as sixty archegonia (Arnoldi and Lawson) in one megaspore; these occur either separately or in some parts of the prothallus they may form groups as in theCupressineae; they are scattered through the prothallus instead of being confined to the apical region as in the majority of conifers. Similarly in theAraucarieaeand inWiddringtoniathe archegonia are numerous and scattered and often sunk in the prothallus tissue. InLibocedrus decurrens(Cupressineae) Lawson describes the archegonia as varying in number from 6 to 24 (Annals of Botanyxxi., 1907). An archegonium consists of a large oval egg-cell surmounted by a short neck composed of one or more tiers of cells, six to eight cells in each tier. Before fertilization the nucleus of the egg-cell divides and cuts off a ventral canal-cell; this cell may represent a second egg-cell. The egg-cells of the archegonia may be in lateral contact (e.g.Cupressineae) or separated from one another by a few cells of the prothallus, each ovum being immediately surrounded by a layer of cells distinguished by their granular contents and large nuclei. During the development of the egg-cell, food material is transferred from these cells through the pitted wall of the ovum. The tissue at the apex of the megaspore grows slightly above the level of the archegonia, so that the latter come to lie in a shallow depression. In the process of fertilization the two male generative nuclei, accompanied by the pollen-tube nucleus and that of the stalk-cell, pass through an open pit at the apex of the pollen-tube into the protoplasm of the ovum. After fertilization the nucleus of the egg divides, the first stages of karyokinesis being apparent even before complete fusion of the male and female nuclei has occurred. The result of this is the production of four nuclei, which eventually take up a position at the bottom of the ovum and become separated from one another by vertical cell-walls; these nuclei divide again, and finally three tiers of cells are produced, four in each tier. In theAbietineaethe cells of the middle tier elongate and push the lowest tier deeper into the endosperm; the cells of the bottom tier may remain in lateral contact and produce together one embryo, or they may separate (Pinus,Juniperus, &c.) and form four potential embryos. The ripe albuminous seed contains a single embryo with two or more cotyledons. The seeds of many conifers are provided with large thin wings, consisting in some genera (e.g.Pinus) of the upper cell-layers of the seminiferous scale, which have become detached and, in some cases, adhere loosely to the seed as a thin membrane; the loose attachment may be of use to the seeds when they are blown against the branches of trees, in enabling them to fall away from the wing and drop to the ground. The seeds of some genera depend on animals for dispersal, the carpellary scale (Microcachrys) or the outer integument being brightly coloured and attractive. In someAbietineae(e.g.PinusandPicea)—in which the cone-scales persist for some time after the seeds are ripe—the cones hang down and so facilitate the fall of the seeds; inCedrus,AraucariaandAbiesthe scales become detached and fall with the seeds, leaving the bare vertical axis of the cone on the tree. In all cases, except some species ofAraucaria(sect.Colymbea) the germination is epigean. The seedling plants of some Conifers (e.g.Araucaria imbricata) are characterized by a carrot-shaped hypocotyl, which doubtless serves as a food-reservoir.
The roots of many conifers possess a narrow band of primary xylem-tracheids with a group of narrow spiral protoxylem-elements at each end (diarch). A striking feature in the roots of several genera, excluding theAbietineae, is the occurrenceAnatomy.of thick and somewhat irregular bands of thickening on the cell-walls of the cortical layer next to the endodermis. These bands, which may serve to strengthen the central cylinder, have been compared with the netting surrounding the delicate wall of an inflated balloon. It is not always easy to distinguish a root from a stem; in some cases (e.g.Sequoia) the primary tetrarch structure is easily identified in the centre of an old root, but in other cases the primary elements are very difficult to recognize. The sudden termination of the secondary tracheids against the pith-cells may afford evidence of root-structure as distinct from stem-structure, in which the radial rows of secondary tracheids pass into the irregularly-arranged primary elements next the pith. The annual rings in a root are often less clearly marked than in the stem, and the xylem-elements are frequently larger and thinner. The primary vascular bundles in a young conifer stem are collateral, and, like those of a Dicotyledon, they are arranged in a circle round a central pith and enclosed by a common endodermis. It is in the nature of the secondary xylem that the Coniferales are most readily distinguished from the Dicotyledons and Cycadaceae; the wood is homogeneous in structure, consisting almost entirely of tracheids with circular or polygonal bordered pits on the radial walls, more particularly in the late summer wood. In many genera xylem-parenchyma is present, but never in great abundance. A few Dicotyledons,e.g.Drimys(Magnoliaceae) closely resemble conifers in the homogeneous character of the wood, but in most cases the presence of large spring vessels, wood-fibres and abundant parenchyma affords an obvious distinguishing feature.